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The genus Cybianthus (Myrsinaceae) in Ecuador and Pern 

John J. Pipolylll 


New flowering plants from southern New Ireland, Papua New Guinea 

W. Takeuchi andjohnj. Pipolylll 


Three new species of Crataegus (Rosaceae) from western North America: 
C. okennonii, C. okanaganensis and C. phippsii 

J.B. Phipps and R.J. 'Kennon 

Taxonomic clarification of Atripkx nxit^llii (Chenopodiaceae) and its near relatives 

Howard C. Stutz and Stewart C. Sanderson 


Flavonoids in Strophostyles species and the related genus Dollchopsis 
(Phaseolinae, Fabaceae): Distribution and phylogenetic significance 
Juan Pablo Pelotto and Maria A. Del Pero Martinez 

Ruellia jimnlcensis (Acanthaceae), a new species from the Chihuahuan 
Desert area, Mexico 
Jose A. VillarrealQ. 


Una nueva especie de Agave, subgenero Littaea (Agavaceae) de Guerrero y 

Oaxaca, Mexico 

Abisai Garcia-Mendoza and Esteban Martinez Salas 


A new species of Mandevilla (Apocynaceae) from Jalisco, Mexico 

Justin Williams 

231 ^^ ^^^K 

A new combination in Mexican Mandevilla (Apocynaceae) 

Justin Williams 


Trachypogon mayaensis (Poaceae: Andropogoneae): A new speaes from Belize 
Joseph K. Wipffand Stanley D.Jones 


Taxonomy of the Sporobolus floridanus complex (Poaceae: Sporobolinae) 
Alan S. Weakley and Paul M. Peterson 


Caryopsis morphology of Leptochloa sensu iato (Poaceae, Chloridoideae) 

Neil Snow 


Xanthostemon fruticosus (Myrtaceae), a new species from the Philippines 

Peter G. Wilson and Leonardo L. Co 

(continued on back cover) 


Volume 18 

Number I 

October, 1998 


Contributions to Botany 


Lloyd H. Shinners 


Wm. F. Mahler 
Publisher 1971-1992 

Barney L, Lipscomb 


Botanical Research Institute of Texas, Inc. 

509 Pecan Street 

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John W. Thieret Prof. Dr. Felix Llamas 

Associate Editor Contributing Spanish Editor 

Biological Sciences Dept. Dpto. de Botanica, Facultad de Biologia 
Northern Kentucky University Universidad de Leon 

Highland Heights, Kentucky 41076, USA E-24071 Leon, Spain 

Guidelines for contributors are available upon request 

and on the inside back cover of the last issue of each volume. 

Subscription per year: $25. Individual, $45. USA Institutions, $50. Outside USA; 

numbers issued twice a year 

®SiDA, Contributions to Botany; volume i8, Number i, pages 1-376. 

Copyright 1998 

Botanical Research Institute of Texas, Inc. 

Printed in the United States of America 

ISSN 0036-1488 



Botanical Research Institute of Texas 

309 Pecan Street 

Fort Worth. TX 76102-4060. U.S.A. 

jpipoly@brit. org: 


The genus Cybianthus was revised to provide taxonomic treatments of tlie Myrsinaceae 
for Flora of Peru, Catalogue of the Vascular Plants of Ecuador, and flora of Ecuador. Eight of 
Cybianthus ten subgenera are represented, and an updated description of the genus, keys 
to its subgenera and emended descriptions for each are provided. Detailed descriptions of 
the morphology, anatomy and ecology of the genus are presented. Cybianthi/s subgenus Iteoides 
is relegated to synonymy under subgenus Microconomorpha . Within each subgenus, keys, 
full descriptions, synonymy, distribution, ecology and conservation statuses, local names 
and uses are given for each species. In addition, specimens are cited for each species, in- 
cluding extralimital ones to show extremes in morphological variation. Thirteen species 
are relegated to synonymy, nine names are lector)'pified, and one, Conoino^fha rigida, is neotypified. 
One new combination, Cybianthus guyanensis suhsp. pse//doicacoreus, is made, and Cybianth/ts 
poeppigit is transferred from subgenus Cybianthus to subgenus Weigeltia. The following 15 
new species are described, illustrated and their phylogenetic relationships are discussed: 
Cybianthus anthuriophyllus. C. cenepensis, C. coniperuvianus. C. croatii. C. flavovirens, C. fosteri, 
C. grandezii. C. granulosus. C. huampamiensis. C. incognitus, C. jensonii. C. nestorii. C. pseudolongifolius, 
C. timanae. C. vasquezii. 


Al preparar tratamientos taxonomicos sobre la familia IMyrsinaceae para los proyectos 
Flora del Peru. Catdlago de las Plantas Vasculares del Ecuador, y Flora del Ecuador, se llevo a 
cabo una revision del genero Cybianthus. Se encuentra en la region ocho de los diez subgeneros, 
y se proveen tanto una descripcion actualizada para el genero como para cada subgenero 
tambien. Se presenta descripciones detalladas tratando la morfologia, anatomi'a y ecologfa 
del genero. Se relega Cybianthus subgenero Iteoides a la sinonimia bajo subgenero Microconomorpha, 
y se transfiere Cybianthus poeppigii del subgenero Cybianthus al subgenero Weigeltia. Para 
cada subgenero, se proveen claves, descripciones completas, sinonimia, distribucion geografica 
y estado actual de su conservacion, nombres locales y usos. Tambien se cita colecciones revisadas 
para cada especie, incluyendo ellas fuera de la region delimitada cuando se muestran variacion 
morfologica significativa. Se relegan 13 especies a la sinonimia, se lectotipifica 9 binomiales 
y uno, Cononwrpha rigida, se neotipifica. Se publica la nueva combinacon, Cybianthus guyanensis 
iuhsp. pseudoicacore//s. Se describen, se ilustran y se discuten el parentezco para 15 especies 
nuevas, listadas a continuacion: Cybianthus anthuriophyllus, C. cenepensis. C. coniperuvianus, 
C. croatii, C. flavovirens. C. fosteri, C. grandezii, C. granulosus, C. huampamiensis, C. incognitus. 
C. jensonii, C. nestorii, C. pseudolongifolius, C. timanae, C. vasquezii. 

SiDA 18(1): 1-160. 1998 

2 SiDA 18(1) 


The genus Cybianthus Marti us was cladistically defined by the presence 
ot unique glandular granules at the junction of the corolla tube and lobes 
by Pipoly (1987). In addition, the unique combination of lateral (axillary) 
inflorescences, gamosepalous and gamopetalous flowers, and stamens con- 
nate by their filaments at least one-fourth their length, and adnate to the 
corolla rube at least one-third its length (Pipoly 19<S7, 1992a) allows for 
practical recognition from herbarium specimens. With this contribution, 
Cybianthus now contains 167 species in ten subgenera. The genus includes 
species formerly included in the genera Comomyrsnn: Hook, f, Couontorpha 
A. DC, Correlliana D'Arcy, Cyh'utnthopin Lundell, GraninuukMict Benth., 
Mkroconomorpha Lundell, and Wetgeltui A. DC. (Pipoly 1 987, 1 992a). In Ecuador 
and Peru, 43 species in 8 subgenera are known. Among the species, none 
are endemic to Ecuador, while 1 1 are endemic to Peru. 

The broad generic concept for Cyhianthi/s was first proposed by Agostini 
(1970) as a precursor to his dissertation (1972); that was followed by the 
formal transfers and a key to the subgenera (Agostini 1 980). Subsequently, 
while revising subgenus Gra)ui>iadenia, Pipoly (1987) determined that the 
paleotropical genus Embelia Burman f. was the sister group to the entire 
genus and that it was most parsimonius to include Gramviadenia within 
Cybianthus. Preparation of taxonomic treatments for the Myrsinaceae in Flora 
of Per//, Catalog//e of the Vasc//lar Plants of Ec//ador and flora oj Ec//aclor re- 
vealed that many taxa were new, and much confusion had arisen among the 
concepts prevalent for taxa already described. Among adjacent countries 
with significant Amazonian regions, Ecuador and Peru share more species 
oi Cybianthus in common than do any other pair of countries. While it would 
have been desirable to include Colombia to provide a treatment for the entire 
northwestern South America, the number of additional species endemic to 
Colombia, or known only from Colombia and Venezuela, would have in- 
creased the size of the treatment by fifty percent. Given that it will be some 
time until a revision of the entire genus for Flora Neotrop/ca is complete, the 
present treatment is mtended to serve as a precursor to that monograph 
and to make the names available for the other aforementioned projects. A 
revised description of the genus Cybianth//s follows, including keys and emended 
descriptions for the eight subgenera occurring in Ecuador and Peru, along 
with keys to the taxa and full descriptions for each. For each of the fifteen 
new species described, illustrations are provided, while for all species, full 
descriptions and synonymy, discussions of distribution, ecology and con- 
servation status, etymology, and when known, local names and uses are elucidated. 


Morphological terms in this treatment follow Lindley ( 1 848) and Pipoly 

PiPOLY, Cybianthus in Ecuador and Peru 3 

(1987, 1992a) for the inflorescence, rachis pedicels and floral parts. Description 
of leaf morphology follows Hickey (1984), trichome description follows Theobald 
et al. (1984) and basic cell and tissue terminology follow Metcalfe (1984). 

Habit and Architecture 

The majority o^ Cybianthus species in Ecuador and Peru are large shrubs 
or small trees to 18 meters tall. Four species are small shrubs or subshrubs 
{Cybianthus croatii. C. huniilis. C. lineatus. C. nanayensis) usually under one 
meter tall. With the exception of two species in subgenus Grammadenia, all 
species are terrestrial. Cybianthm marginatum is an obligate epiphyte, while 
C. magnus is a facultative epiphyte, frequently found on large tree trunks, 
but also known from large rock outcrops. 

Most members g{ Cybianthus exhibit Rauh's Model of architecture (Halle 
et al. 1978), characterized by a polyaxial, monopodial, rhythmically grow- 
ing, readily distinguishable trunk, that develops tiers of branches morpho- 
genetically identical to itself. All branches are orthotropic and monopo- 
dial, with lateral (axillary) inflorescences that do not affect shoot development. 
However, two subgenera {Comomyrsine and Triadophora) exhibit Corner's Model 
(Halle et al. 1978), characterized by a single aerial meristem that produces 
a monoaxial (unbranched) axis on which inflorescences are lateral (axillary). 
The resulting monocaulous tree is pleionanthic (not hapaxanthic, or mono- 
carpic) and growth is indeterminate. The trunk may grow rhythmically or 
continuously. Sporadic occurences of this architectural model occur in members 
of subgenus Weigeltia, from the Guianas and eastern Amazonia, however, 
none of those species are known from Ecuador and Peru. 

While conducting tieldwork in Peru, a collection o1 Cybianthus kayapii 
(Lundell) Pipoly was observed reiterating. In the population observed at 
Explorama Lodge near Yanamono, (Loreto, Peru) one individual {Pipoly et 
al. 12383), exhibited bayonet reiteration, caused by destruction of the api- 
cal meristem of the trunk. The individual bore a reduced staminate inflorescence 
with flowers slightly larger and more maroon in color than normal, and smaller 
leaves with shorter petioles. As I have noted elsewhere (Pipoly 1987, 1992a), 
leaves on the reiterative shoots resemble those of saplings. Variation in 
inflorescence and floral morphology seen in this Peruvian population is consistent 
with variation reported in Pipoly (1992a). Therefore, it appears that changes 
in sex expression are consequent to a reiteration phenomenon, rather than 
circumstantially associated with it. I postulate that reiterative branches may 
be hormonally juvenile and iis such, are morphogenetically incapable of producing 
bisexual or pistillate flowers, and produce only sapling leaves until flowering 
occurs. This process may account for the great variation in leaf morphology 
and apparent sexual lability often associated with many species of Cybianthus. 
Unfortunately, for individuals of normally dioecious species with monoaxial 

4 Si DA 1<S(1) 

models, it is not possible to determine if sex change has occurred because 
no inflorescences are usually observed before the original apical meristem 
is destroyed, and no inflorescences on other branches are available for com- 
paris{)n. Hov^^ever, precociously flowering individuals o'iCybtanthiis incognitiis 
(P. Barbour 2403) stipport the hypothesis. Clearly, pruning experiments /'// 
.r//// will be necessary to test this hypothesis. 


A branchiet is here defined as the distal 10 cm oi any branch. Branchlets 
may be flexuous, or straight; terete or angular; smooth, verructflose, or ver- 
rucose-papillate (Fig. lA, IB), glabrous, glandular-papillate (Figs. ID, 3F, 
4F), ferrugineous stipitate-lepidote (Fig. 2A— F), ferrugineous or rufous dendroid- 
and/or stellate-tomentose (Fig. 3A), rufous sessile furfuraceous lepidote, with 
a tomentum of malpighiaceous trichomes (Fig. 3D), glandular-granulose, 
with hydropotes (Fig. 3C), epunctate or rubiginous punctate-lineate, bear- 
ing lenticels or not. 


Species ofCyi?/a}ith//.s\ like all Myrsinaceae, are exstipulate and have simple 
leaves. The leaves are usually alternate, but may be subopposite or 
[:)seudoverticillate, especially in subgenera. Mkroaj/wf/mpha, Co//umyrs/m, Weigeltia 
and Cybianthus . In the majority of species, the leaves are petiolate, but in 
subgentis Grammadenia they are sessile. The ptyxis (the form in which one 
shigle leaf is folded on itself in btid) is most often supervolute (Cullenl978), 
but it has not been adecjuately studied thtis fitr for each subgenus. The leaf 
vernation sensu CuUen (1978), is at least analagous with floral aestivation, 
or the relationship of one leaf to another; in Cybianthus, it is in fact, un- 
known. The leaf blade texttire may be membranaceous, chartaceous or co- 
riaceous. The shape is most often elliptic, but may be oblanceolate, lan- 
ceolate, obovate, rarely ovate or oblong {Cybianthus nuirginatus). The apex 
may be acute, acuminate, long-attenuate, rounded, obtuse, or emarginate, 
without mucro or rarely (stibgenera Grammadenia, Comomyrsine, Triadophora) 
mucronate (Fig. 21). The base is rarely auriculate (subgenus Grammdenia) 
or obtuse approaching auriculate {Cybianthus grandezii. C. kayapii. C. 
anthuriophyUus) and is usually cuneate, long-attenuate, acute or obtuse, decurrent 
on the petiole or not. The adaxial surface may be nitid, pallid, or sordid, 
rarely scrobiculate , pusticulate or pustulate when mature. The blade mar- 
gin may be regular or irregular, flat inroUed or revolute, entire, or rarely 
undulate, lobate, crenate or dentate {Cybianthus pastaisis), densely and mi- 
nutely serrulate {C, anthuriophyUus), or coarsely dentate {C. schlimii. some 
populations of C pastensis, C. poeppigii, ), or with a very subtle btit vascular- 
ized vein ending (C verticiHoides) opaque, or rarely scariotis (all species in 

PiPOLY, Cybianthus in Ecuador and Peru 

Fig. 1 . SEM photos ot morphological features in subgenus Grammadenia. A. Cybianthus marginatus 
{'Pipoly 6546), showing verrucose papillae. B. Same, close-up. C, D. C. lineatus (Pipoly et al. 
7729), showing stem surface. D. Glandular papillae. E. C. costarkanris subsp. morii {Pipoly 
7017), pollen, equatorial ant! polar views. (Bars in figs, equal: A. 50 pm, B. 22.2 pm, C, D. 
48 pm, E. 12.3 pm) Figure reproduced from Pipoly, 19H7. 


SlDA US(1) 

Fk;. 2. SEM photos of representative stipitacc fcrrugincous lepiclote scales of SLibt;enus Ciiniiiindrlihii. 
A. Cyhiduthiis {liii^autiiphylh/s (holocyjie). V>. C. spichii^^eri (holotype). C. C. hittiis (D/nlley lOHOi). 
D. C. per/n-hiiii/s (Dodsofi 282 1 ). E. C. comper/iviarms {Killip &. Smith 260'^ 3). F. (2 )^/tytiiiciisis 
subsp. p.seiicliiiuiioyei/s {Siiiipsoii & Schinikc 7,S4). (Bars in Figs. A-F ec]ual: 50 pm). 

CybicDitlv/s subgenus Gramnmdema; Cyhianthiis h//»/ilis. and C. troatii ot Cyhuniih/is 
subgenus Conioriiyrsim). 

Leaf punccation and punctate-lineation may be pellucid (translucent), black, 
orange red, or rubiginous (light red) in color. Punctations as defined by Pipoly 
(19H7) are definded as rounded, lyso-schizogenously formed cavities and 
are distinguished from ptmctate-lineations, which are homologous, linear 
cavities in the leaves. Among taxa bearing non-pellucid punctations or punctate- 
lineations, all young, undexpanded leaves have orange glands, even il ma- 
ture leaves have red or black glands. Experiments to determine integrity of 
punctation color have revealed that it is not a reliable taxonomic character 

PiPOLY, Cybianrhus in Ecuador and Peru 

0t "* 

Fig. 3. SEM photos ot representative vestiture in subi2,enera Laxiflorus. Mkrocojioinorpha. Triadophora 
and Coiitoinyrsine. A. Cyh'uinthin fiilvopiilvtviikntus stibsp. magnoliifolitis {Cowan & Soderstrom 
2146), branchlet, showing' stellate trichomcs. B. C, sp/cati/s {Maguire & Puliti 28064), showing 
hytlropote in abaxial leaf pit. C. C. pcisttusis (Killip & Garcia 33886), branchlet, showing 
developing hydropotes. D. C. scblinui {Fosherg 20148), branchlet, showing malpighiaceous 
trichome. E. C. kayapii (holocype) Sraminate flower, showing vestigial pistillode, structure 
of androecium. F. C. hitniilh {l.ehmanii s.n.), showing glandular papillae of abaxial leaf sur- 
face. (Bars in Figs. ec]ual: A-C. 50 pm, D & F. 25 pm, E. 0.4 mm), 

(Pipoly 1987). The other punctation character states include "conspicuous" 
versus "inconspicuous," reflecting the ease of viewing the punctation (a reflection 
of its relative diameter in transverse section), and "prominent" versus "not 
prominent," which refer to whether the punctation is blister-like or planar, 

The abaxial and/or adaxial surface of the blade may be glabrous, or with 

8 Si DA 18(1) 

a vestiturc composed of stipirate ferrugineous lepidote scales (subgenus 
Cononm-pha, Fig. 2A-2F), sessile fi.irfuraceous lepidote scales (subgenus Weigeltia), 
dendroid and/or stellate crichomes, translucent glandular-lepidote scales, 
or with scattered glandular papillae (subgenus Coiuoniyrsine, Figs. 3F, 4F ), 
maljMghiaceus trichomes (subgenus Triadophora , Fig. 3D), or hydropotes 
(subgenera Laxiflorns, Microcoiioiiiorpha, Grannnadenia, Coniomynine^ Figs- 3A, 
3C, 4A-E ). The adaxial surface is usually giabrescent. 

Hydropotes ("water drinkers") were previously thought to be unique to 
subgenus Granivuknia (Pipoly 1987), but have now been found in all spe- 
cies of subgenus Comnniyrsine. Described by Mayr (191 5) and Gruss (1927a, 
1927b), hydropotes have been reported for numerous submerged aquatic 
angiosperms (Fahn 1979; Gessner & Volz 1951). Elegant ultrastructural 
and autoradiographic studies undertaken by Liittge (1964) and Liittge and 
Krapf (1972) demonstrated the mineral absorptive function of hydropotes. 
While both subgenera bear these structures, their morphology is strikingly 
different. The hydropotes of subgenus Grammdenia (Fig. 5) consist of five 
to seven subsidiary cells, a central foot cell, a basal cell, a stalk cell, and up 
to 12 cells forming a scale-like cap, while those of subgenus Comniiiyrsine 
consist of five to seven subsidiary cells, a central foot cell, a stalk cell, and 
up to 1 2 cells forming a lenticular cap or globose body. In subgenus Conmiiyrsme, 
the cap cell is formed before elongation of the stalk cell. In Cybianthus verticilloides 
(Fig. 4B), C. spruce! (Fig. 4D), and C. kayapii (Fig. 4E) the cap consists of a 
spherical body of cells, while m C. croatii (Fig. 4 A) and C, simplex (Fig. 4C), 
the cap is lenticular. It is interesting to note that within subgenus GraniDuideuui, 
the shape of the caj-) is the same among all species, while in Comimiyrsiue^ 
there are 4 types. The occurrence of hydropotes in subgenus Laxijior/zs, in 
pits over the abaxial leaf surface, and in subgenus Microconomorpha, under 
the papillate tomentum of the branchlets, was unknown before the present 
study. Despite these differences, the structures are homologous, based on 
position, function and ontogeny. In early ontogeny, a mucilaginous sub- 
stance accumulates around the base cell (Fig. 7E). Later, the cap breaks off 
(Fig. 7F), leaving a mucilaginous ring around the broken basal cell (Pipoly 
1987, unpubl. data). 

Subepidermal fibers, visible as numerous, thin, parallel lineate structures 
on both surfaces of leaf blades in subgenus Triadophora are unique to it. They 
have recently been shown to be homologous with those of Clavija in the 
Theophrastaceae (Pipoly, unpubl.). 

Petioles are present in most taxa, with the notable exception of subge- 
nus Gram/i/deiiia, where the leaves are sessile. The petioles may be canaliculate, 
marginate, or canaliculate with margins, trigonal, or rarely subterete, and 
may be pulvinate (abruptly swollen basally) or gradtially tapering basipetally. 

PiPOLY, Cybiantlius in Ecuador and Peru 

Fig. 4. Representative SEM photos ofmorphological features in subgenus Comouiyniut. A- 
E, Hydropotes of abaxial leaf surface. A. Cyhianthns croatii (holotype), showing lenticular 
cap. B. C. vertidUoides (holotype), showing globose cap. C. C. iimplex {Zak 1350), showing 
lenticular cap. D. C. spruce/ (C//atrea/sas 15743), showing globose cap. E. C. kayapii (holo- 
type), showing depressed-globose cap (somewhat collapsed). F. C. kayapii (holotype), branchlet, 
showing glandular papilla. (Bars in Figs. ec]ual: A. 25 pm. B. 20 pm. C. 10 pm. D— F, 25 

Cataphylls and Pseudocataphylls 

Cataphylls (Fig. 9B) and pseudocataphylls are synapomorphic to subgenera 
C omomyrs ine 3.nd Triadophora, respectively. Earlier (Pipoly 1987), I had mis- 
takenly interpreted them as inflorescence bracts. They may be alternate or 
pseudoverticillate, alternating with pseudoverticels of leaves, or apparently 
axillary to them, rigid, chartaceous, or membranaceous, linear-subulate to 


Si DA IcSd 

V]i,. 5 . Onrogeny ot hydroporcs in siilit;c-nLis GrciiiiiiUHk)iui. A, B, & D. Cyhuintbiis aistciyiunim 
subsp. cintciricciHiii {Pipfily 76(AS'), SHM photos. A. MatLire hyclropore, cap with at least 8 
cells, anti with 5 subsidiary cells. B. Younger hydropote, cap with 1 cells, ami with 6 sub- 
sidiary cells. C. C CDSliiyncimis stibsp. iiinri'i (Pipiily 701 7), longisection of yoting hydropote, 
showing sLibsidiary cells (sc), loot cell (fc), stalk cell (st), cap cell (ca), basal cell not dis- 
cernible. D. Young hydro|-iote, with 2-celled cap, showing miicilagmous SLibstance (ms). 
E, K C. coitarkaniis subsp. [hii/iiiiifiisis (Pipuly 7036), paradermal sections. E. Showing ma- 
ture hydropote, with 1 2-celled-cap. V. Showing hydropote after cap has broken off. (Bars 
in Figs. A-F equal: A & C 28 pm, B. 20 |_im, D. 10 pm, E. 36 pm, F. 5.i pm. Figure 
reprodticcd from Pipoly, 1987. 

PiPOLY, Cybianthus in Ecuador and Peru 11 

acicLilar, rigid to membranous, keeled or flat, apically mucronate or not, 
prominently punctate or punctate-lineate, glabrom, aifous pubemlent, glandular- 
papillate, bearing hydropotes, or orange lepidote scales, sessile or on a peti- 
ole to 2 mm long. Pending further developmental studies, I am distinguishing 
cataphylls from pseudocataphylls based on the absence of a petiole in the 
former and its presence in the latter. Within Cybianthus, pseudocataphylls 
are unique to subgenus Triadophora, while cataphylls are unique to subge- 
nus Comomyrsine. Both pseudocataphylls and cataphylls may be distinguished 
from an inflorescence bract by the fact that neither of these structures are 
axillant to an inflorescence, neither are ephemeral, but both occur in pseudowhorls. 

Inflorescence and Flowers 

The inflorescence in Cybianthus is always lateral (axillary), and it may consist 
of a simple raceme (erect or kix), a subpyramidal or pyramidal panicle (sometimes 
thyrsoid), a spike, or rarely, a pleiochasium. At times, species with essen- 
tially racemose inflorescences may produce a panicle consisting of a pseudoverticel 
of racemes on a reduced peduncle. In this treatment, an inflorescence is considered 
spicate if the pedicels are obsolete or less than 0.3 mm long and subspicate 
if the pedicels are from 0.4-0.8 mm long. In most species, the staminate 
inflorescence tends to be longer, slightly more lax, and bears greater num- 
bers of flowers. In those species bearing panicles, the staminate ones bear 
secondary branches of the inflorescence that are longer, more floriferous, and 
at times, more branched than the pistillate and bisexual panicles. 

The inflorescence bracts are early caducous and are rarely seen in the field 
or on specimens. The secondary branch bracts of panicles are also rarely collected, 
with the exceptions of subgenera Comomyrsine and Triadophora. The floral 
bracts may be membranaceous or chartaceous and are usually persistent in 
staminate inflorescences, but are at times caducous in pistillate ones. The 
floral bracts may be glabrous or bear a tomentum either adaxially, abaxially, 
or on both surfaces. The floral bract apices are acute, attenuate, or acicular, 
the margins entire or undulate. 

The pedicels are cylindrical, clavate, or obconic, and at times accrescent 
or crassate in fruit. In most species, the pedicels are subtended by an axillant 
floral bract, but in Cybianthus kayapii, it is inserted at about the middle of 
the pedicel. 

Figure 6 illustrates the tremendous variation in floral morphology among 
members of the genus, along with representative staminate and pistillate 
flowers from Embelia, the paleotropical sister genus o{ Cybianthus. The flowers 
may be functionally staminate, pistillate or bisexual. Consequently, the plants 
are normally functionally dioecious, but may also be bisexual or polyga- 
mous. Flowers are normally perfect, but in some species of subgenera Weigeltia, 
Comomyrsine and Cybianthus, the pistillode is often obsolete in the stami- 




il}{::^^;.-- .:^W] 


p. wVi 

1 mm 

Fi(i. 6. Rc-|Tr(.-sc-nranvf flowers of C)'/'/f/«//v/.f subgenera antl sister genus, £w/W/;/, open flowers 
in anriiesis above, calyx lobes below. A-B, Subgenus A\/cri)aji/o///orpha, {Cy/?/ai/fh/fs /),/.\-/ei/s/s), 
note monomorphic flowers. A. Srammate flower {Killiji & Cuira'ct }}SS(^). B. Piscillare flower 

PiPOLY, Cybianthus in Ecuador and Peru 13 

nate flowers. The staminate flowers are usually larger than the pistillate ones. 
The flowers are usually 4 or 5(— 6)-merous, but they are 3-merous in subge- 
nus Triadophora (Fig. 6K, 6L). The majority of species are homomerous, but 
Cybianthus kayapii has heteromerous flowers, (the calyx 4- and the corolla 
5-merous), and in C. anthuriophyllus, the calyx 6- and the corolla 5-merous. 
Either or both whorls of the perianth may be membranaceous, chartaceous, 
coriaceous or carnose, epunctate, punctate or punctate-lineate. The punc- 
tations may be inconspicuous or conspicuous, prominent (raised) or not, 
pellucid, brown, orange or red. The calyx may be valvate or imbricate, and 
may be cotyliform, cupuliform or urceolate. The corolla is valvate or im- 
bricate, and may be campanulate, cupuliform, tubiform, infundibuliform, 
salverform, rotate or subrotate. The stamens and staminodes are similar in 
morphology, but the staminodes are smaller. The filaments of the stamens 
and staminodes are partially united at the base to form a conspicuous or 
inconspicuous, membranaceous, chartaceous or carnose tube, except in subgenus 
Cybianthus, which has a terete or rarely, angulate staminal tube develop- 
mentally fused with the corolla tube, the stamens thus appearing epipetalous. 
In some species, the tube bears lobes (sterile projections of tissue) which 
alternate with the apically free portion of the filaments. The apically free 
portions of the filaments may be terete, flat, or absent. The anthers may be 
dorsifixed, basifixed or versatile. Anthers may be lanceolate, ovate, cuadrate, 
or deltate, apiculate, acute, truncate or emarginate, the apiculate ones may 
have the apiculum erect, distally or proximally recurved, glabrous or glan- 
dular-papillate. Antherodes of the pistillate flowers are similar to the an- 
thers but reduced insize, and normally devoid of pollen, but occasionally, 
they may produce copious amounts of abortive pollen (Pipoly 1983a). The 
connective may be epunctate or prominenrly punctate, or rarely glandular- 
papillate. The pollen is tricolporate and psilate (Fig. IE). The pistillode 
may be lageniform, conic or obturbinate. The pistil may be obnapiform, 

{Schidtes 3226). C-D, Subgenus Stapfia, {Cybianthus stapjii). C. Staminate flower {Fassett 25 706). 
D. Pistillate flower {Cuatrecasas 1229.5). E-F, Subgenus Laxiflori/s {Cybianthus spicatus). E. 
Staminate flower (Al^^/^'/rf 33523). F. Pistillate ^ov'/ts: {Prance et al. 3339). G-H, Subgenus 
Conomorpha {Cybianthus laurifoUus) . G. Staminate flower {Garcia-Barriga &Jaramillo 19841). 
H. Pistillate flower {Grubb etal. 744). I-J, Subgenus Comomyrsine. L Staminate flower, Cybianthus 
sprucei {Cuatrecasas 1 6272). J. Pistillate flower, Cybianthus kayapii {Acosta-Solis 5544). K-L, 
Subgenus Triadophora, (Cybianthus schlimii). K. Staminate flower {Haught 1502). L. Pistil- 
late flower (Herrera 1 720). JM-N, Subgenus Weige/tia, Cybianthus sp. nov. iVI. Staminate flower 
{Maas & Westra 4459). N. Pistillate flower {Maguire & fansljawe 405 90). O. Subgenus Cybianthopsis, 
bisexual flower, {Cybianthus sintenisii, P. Wilson s.n.). P. Subgenus Grammadenia, bisexual 
flower, {Cybianthus piresii, Maguire et al. 37052). Q-R, Subgenus Cybianthus, {Cybianthus 
goyazensis). Q. Staminate flower {Irwin & Soderstrom 5378). R. Pistillate flower {Herringer 
45). S-T. Embelia sp. S. Staminate flower {Petelot 3887). T. Pistillate flower (C. Wang 76408). 
Drawing prepared by Peggy Duke. 

14 SiDA 18(1) 

conic, ellipsoid, or obturbinate, with a punctiform, or capitate stigma. The 
stigma is large and capitate, with erose-fimbriate lobes and is caducous in 
subgenus Coniomyrslne. The ovary in species from Ecuador and Peru are ter- 
ete. The placenta may be cotyliform, cupuliform or globose, with 2—4 uniseriate 
ovules naked or partially immersed. The fruit is a globose or depressed-globose 
drtipe and is one-seeded, with a thick or thin exocarp. 


This treatment is not intended to serve as a monograph of the genus, but 
a few salient anatomical features may prove useful in identifying sterile material 
or wood samples. Druses (Pipoly 1987-Fig. 8B— C) are ubiquitous in iVLyrsinaceae, 
as are pericyclic fibers (Pipoly 1987, Fig. 8A). All Myrsinaceae have resin 
ducts (canals) at least in the cortex, and in the field, copious amounts of 
resin are visible in the canals of the pith and secondary phloem, in freshly 
cut branchlets, in species of subgenera Grctuimadenta and Laxiflorns. Aeren- 
chyma in the cortex of the primary stem is unique to subgenus Grammdenia 
and is found in all species (Fig. 7F), except in Cybianthns lineatm (Fig. 7E). 
The pith in primary stems of subgenus Weigeltia is parenchymatotis with 
large, rcninded starch grains, while that of Cyhianthiis ))icigniis has angular 
coUenchyma (Pipoly 1987). It is interesting to note that the collencyma of 
the outer cortex m CyhuinthHS magniis subsp. asynimctrici/s is tangential rather 
than angular, while that of C. inapun subsp. magnns is angular (Fig. 8 D— 
F). All species of Cybianthiis have cortical vascular bundles, that may be 
amphicribal (Fig. 7C) or hemiamphicribal (Fig. 7D), with (Fig. 7C) or without 
(Fig. 7D) accompanying perivascular fibers. Wood of subgenus Comoniyrsine 
is notable for its thin-walled vessels (Fig. 9D), while the fiber-tracheids of 
subgenus Grcimmadenui have walls so thick as to significantly occlude the 
lumina (Pipoly 1987- Fig. 7C). Also, starch is present in the phloem fibers 
of subgenus Comoniynnn: (Fig. 9F, 9F) but not in Graninukuut. In leaf anatomy, 
it is notable that subgenus Grcn)Wicule>iui has functionally acrodromous ve- 
nation. This is due to its unique leaf-node continuum, and consequent pri- 
mary vascular system (Pipoly 1987-Figs. 7-9; 1 1-12) where the cathodic 
and anodic leaf traces are autonomous from each other and from the rel- 
evant midrib trace, a system thus far unique among angiosperms. Leaves of 
subgenus Tnadophora may be easily recognized by their subepidermal fibers, 
while those o'iCybicnithiis Inimtits are unicjue for their bifacial palisade layer 
(Pipoly 1987- Fig. 14b). 


In Ecuador and Peru, species of Cyb'nnith/is occur in wet or moist, tall 
terra firme foKst on laterite, limestone or white sand, seasonally inundated 
igapo or varzea, premontane humid, wet or pluvial forest on laterite or sandstone. 

PiPOi.Y, Cybianthus in Ecuador and Peru 


Fk;. 7. Primary stem iiistological featLires in subgenus Gninnnackiiid . A— B. Cybianthus ptariensis, 
(Pipoiy et al. 7133), periderm formation, showing epidermal cork development. C. C. marginatum 
{Pipoly 6346), amphicribal corical bundle without periv iscular fibers. D. C. ptariensis {Pipoiy 
et al. 7133), hemiampliicribal bundle with perivascu'ar fibers (pf). E. C, limatiis {Pipoiy 
7229), section showing parenchymatous pith, mnct and outer cortex. F. C. marginatiis (Pipoiy 
6546), section showing aerenchymatous mner cortex; parenchymatous pith and outer cor- 
tex. (Bars in Figs. A-F equal to: A, 30 [.mi, B. 1 20 pm, C. 48 pm, D. 60 pm, E. 465 pm, 
F. 120 pm. Figure reproduced from Pi|-)oly, \'-)H7. 

cloud forest, elfin forest, montane or subparamo thickets, or sandstone scrub 
at high elevation. 

The terra firme and premontane forest life zones are the richest in endemics, 
with six and five species, respectively. Terra firme is here divided into two 


SiDA 18(1) 

Fig. 8. Primary stem histological leacures of subgenus Grammadenia . A. C. costarkanm subsp. 
morit {Pipoly 7017), showing pericyclic fibers (arrow). B. C. costaricat/j/s subsp. costaricanus 
{Pipoly 7068), showing druses in pith. C. C. costaricanus suhsp. pa>2ai//ens/s {Pipoly 7056), 
showing druses in pith, using polarized light. D, E. C. magniis %\}i0^p.magnus {Pipoly 6453), 
showing angular pith collenchyma. F. C. magnits subsp. asyiiimetricus {huteyn, Pipoly et al. 
10415), showing transitional tangential pith collenchyma. (Bars in Figs. A-F equal to: A. 
195 pm, B. 45 pm, C. 48 pm, D. 230 pm, E. 74 pm, F. 60 pm.) Figure reproduced from 
Pipoly, 1987. 

edaphic categories, lateritic and white sand (often referred to as "variUal" 
by Peruvians) and is defined as lowland tall moist or wet forest which is not 
inundated. The canopy normally reaches .35 meters, with very few emer- 
gent individuals. In the lateritic terra firme, Cybianthm kayapti, C. schlhnn, 
C. pseuJoIong/folius, C. vasquezii, C. cenepensis, C. grandezh, C. fuscus, C. jemonii, 

PiPOLY, Cybianthus in Ecuador and Peru 


y E 



Fig. 9. Morphological and anatomical features of subgenus Coniornyrsine {Cybianthus kayapii, 
Pjpoly et al. 12490). A. Habit, showing bayonet reiteration. B. Habit, showing pyramidal, 
bipinnate panicle and leaf tapering to obtuse base. C. Stem apex, showing cataphylls. D. 
Transverse wood section, showing thin-walled vessels. E. Tangential wood section, show- 
ing rays and phloem fibers-tracheids. F. Phloem fiber-tracheids, showing starch grains. (Bars 
in Figs. A-F, A. 12.6 cm, B. 2 cm, C. 1 cm, D. 400 pm, E. 100 pm, 

18 SiDA 18(1) 

and C. venez/ielanus arti found. Cyhianthm kayapii, C. schlifiiii, and C. grandezii 
are found on steep banks of small creekbeds, C. cenepensis and C. venezuelanus 
are ridgetop species, and C. cenepensis, C. jensonii, C. vasqi/ezii and C. pseudolongifolius 
grow along terraces above larger rivers. Vasquez ( 1 997) has described "Irapayal," 
"Yarinal" and "Supay Chacras" associations within the lateritic terra firme, but 
with so few comparative, quantitative forest inventories completed for both 
countries, I am not able to precisely list the Cybianthus species known from 
each. The terra firme forests on white sand (varillal) are shorter in stature than 
those on lateritic soils, and generally support lower numbers of lianas. Among 
the varillal forest types described for Peru, Vasquez (1 997) lists "varillal seco," 
"varillal humedo" and "chamizal" or "ojo de varillal" as.sociations, only some 
of which have been noted on label data. However, Cybianthus perinnanus , C. 
spichigeri, and C. gigantophyllus occur on rolling hills in the varillal forest 
type. Among those species which occur on steep hillsides near light gaps, 
Cybianthus gigantopyllns is most notable, as it occurs in the ecotonal area 
between forest gaps and mature forest. Cybianthus nanayensis, a subshrub, is 
frequently found in gaps left by large treefalls in overmature forests, and 
along trailsides, where it occurs in the rather dense herbaceous stratum. Cybianthus 
resinosus, another inhabitant of the terra finite forest on white sand, occurs on 
terraces above black water rivers in the forest, while C. nestorii is found in 
the more open shrubby "varillal seco" transitional area near the riverbank. 
Cybianthus Julvopulverulentus suhs\^. Diagnoliifolius typically occurs in campina, 
or campinarana formations m Brazil, but in Peru it has been collected once 
in the "varillal seco" an open shrubby area on white sand several hundred 
meters from a black water riverbank. Unfortunately, no literature directly 
addressing this forest type is known for Ecuador. 

The two forest types subject to inundation have been divided mto varzea, 
flooded by white water, and igapo, flooded by black or black and white water. 
The other significant difference between these two forest type is that in varzea, 
the forest is indundated for a much shorter time than that of the igapo. To 
date, I know of no documentation for occurrences of forests inundated by 
clear water in Ecuador or Peru, as they are found elsewhere in Amazonia 
(Brazil, Colombia, Venezuela), but are best developed in Colombia. Three 
species are known from igapo, with no endemics. Cybianthus guyanensis subsp. 
pseudoicacoreus is found in the igapo at its limit with terra firme, while C. 
penduliflorus is found well within the igapo and is frequently found in standing 
water. Cybianthus spicatus is found both in igapo and varzea, and exhibits 
both staminate and pistillate ecotypes, with some exhibiting apparent ran- 
dom variation. It is a broadly ranging polymorphic ochlospecies (sensu Pipoly 
1983a) with great morphological variation. The known Peruvian popula- 
tions, from Huanuco and San Martfn, are identical to those found in cen- 
tral Guyana (Pipoly 1 9H3a). The other varzea species, Cybianthus cydopetalus. 

PiPOLY, Cybianchus in Ecuador and Peru 19 

is known only from Madre de Dios in Peru, and from the Jurua area in Brazil. 
It grows in the margin of varzea near its junction with the terra firme forest. 
Forest associations within the vazea have been described by Vasquez (1997), 
but I have been unable to match the corresponding complement oiCybianthus 
species, owing to inadequate label data. The three forest associations present 
in varzea for Peru include, "barrillal," "restinga" and "bajial." Clearly, more 
fieldwork is needed to discern foristic differences among these association 

Premontane forest habitats are found from scarcely above 200 m to nearly 
1,000 m. The forest is lower in stature than the terra fir me, have a greater 
epiphyte load, and a larger number of lianas. Those on sandstone are dis- 
tinguished here from those on other soils. On sandstone, three species are 
found of which one, Cylnanthus timanae, is endemic. The other species, Cybianthus 
comperuvianus , a new taxon described herein, is known from these forests in 
Peru, Bolivia and adjacent Brazil. The other premontane forest formation 
on lateritic soils hosts 9 species, Cyhtanthm minutiflorm, C. huanipamiemis, 
C. granulosus, C. poepptgii, C. schlmiii, C. peruvianus, C. fosteri, and C. venezuelanus 
and C. flavovirens. Among these, only Cybianthus venezuelanus, C. poeppigii, 
C. schliniit and C peruvianus are not endemic. A surprising new distribu- 
tion record for Cybianthus lepidotus, from Bagua Province, Imaza District, 
of Amazonas Department, Peru, is recorded here. Cybianthus lepidotus was 
once thought to be a Guayana Higland endemic, but was found in Bolivia 
in the Maipiri region on the sandstone "laja" formations there (Pipoly 1992a). 
This kind of disJLinction, concomitant with those for species such as Cybianthus 
spicatus, and Cybianthus lineatus (see below), support recent thematic map 
data at NASA, showing that the Ecuadorean/Peruvian area north, slightly 
east, and immediately south of the Cordillera del Condor, contains significant 
sandstone formations that may constitute "tepui satellites" (sensu Maguire 
1979). Clearly, much more exploration in southeastern Ecuador and north- 
ern Peru, should be a high research priority. 

There are six species oiCybianthus known from cloud forests. Cybianthus 
pastensts and C. incognitus are found in areas of high shade and moisture within 
these forests. Along the margins to the leeward side, in that portion of the 
Choco Floristic Province that extends into Ecuador, Cybianthus cuatrecasasii 
may be found, while on the eastern Andean slopes of Peru, C. laetus grows 
in exposed, wind-swept margins of the forest. The lack of records for Cybianthus 
laetus in Ecuador is more likely a collection artifact than a reflection of its 
rarity, given its occurrence in the Department of Boyaca, Colombia. It is 
notable that Cybianthus patensis and C. incognitus, when growing at eleva- 
tions below 1 ,000 m, are ridgetop species. Cybianthus magnus subsp. ?nagnus 
is an obligate epiphyte in closed cloud forests, growing in the forks of trees 
or on deep detritis, and shielded from winds. 

20 Sum 18(1) 

Elfin forests and montane thickets are transitional formations below the 
jalca ( a formation like a paramo but without species of the Asteraceae subtribe 
Espletiniae). Elfin forests host considerable numbers of trees and usually 
have trees up to 5 m tall. The montane thicket (subparamo) grows in more 
exposed areas at higher elevations and is dominated by low shrubs and small 
trees to 2 m, with broad crowns. Cyhianthus viarginatiis is found in both of 
these habitats, but the leaves and stature of the plants are much smaller in 
the thicket formation. Also, the verrucose papillae of the stem are much 
more pronotmced in the thicket habitats than in plants growing in the elfin 
forest. Cyhianthus, viagnus subsp. asymmetries grows in open montane for- 
ests or subparamo thickets, and elfin forests, where it is subject to exposure 
to the high winds and rain. Its leal variation is significant, but it is easily 
recognized from subspecies magnus by the hydropotes of the adaxial leaf surface 
and orange punctations of leaf, inflorescence and perianth parts and the unique 
white, then lavender fruits. 

Finally, sandstone scrub, called "pajonal," is known thus far only from 
Peru. It is the formation growing at the highest elevations where JVIyrsinaceae 
occur, mostly well over 3,000 m. In these habitats, there are few, small shrubs 
which rarely exceed 1.5 m tall. Cyhianthus I meatus is found in this habitat, 
the first locality for this species outside the contiguous Guayana Highland. 
No similar habitat has been described in Ecuador, but it may be present in 
the Cordillera del Condor. 


JVty species concept follows that of Wiley ( 1 978, 1 98 1 ), who defined a spe- 
cies as follows: "An evolutionary species is a single lineage of ancestor-descen- 
dant populations which maintains its identity from other such lineages and 
which has its own evolutionary tendencies and historical fate." My subspecies 
concept (Pipoly 1987), defines a subspecies as follows: "groups of populations 
within a single lineage of ancestor-descendant populations that show varia- 
tion by unic|ue combinations of plesiomorphies, or homoplasic apomorphies, 
correlated with biogeography and/or ecology. This rank is primarily used to 
convey information regarding variation in the life histories of these populations 
and character state differences hypothesized to be the result of this variation. 
The subspecific rank in no way attempts to predict speciation events." 

The keys are artificial and designed to expedite identification of herbarium 
specimens. An attempt has been made to emphasize vegetative characters to 
increase the keys' usefulness with sterile material. The numbers appearing be- 
fore the taxa refer to their respective position in the key; any correlations with 
phylogenetic relationships are coincidental. Quantitative and cjualitative 
data presented in keys and descriptions for floral parts and bracts were taken 
from organs rehydrated from herbarium specimens by boiling in water. 

PiPOLY, Cybianthus in Ecuador and Peru 21 

Measurements from these range from 10% to 15% greater than those mea- 
surements taken directly from dried material. Data regarding stem diam- 
eters, inflorescence rachises, pedicels, leaf, and fruit shape were taken from 
dried herbarium specimens. Extra-Ecuadorean and -Peruvian specimens are 
cited for all new species and for recent collections of other species used to 
significantly amplify previously pubHshed morphological descriptions (Pipoly 
1981, 1983a, 1983b, 1987, 1988, 1991, 1992a, 1993, 1994, 1995, 1996). 
A description of the genus Cybianthus and a key to its subgenera in "Ecua- 
dor and Peru are provided below. This description, along with that of the 
subgenera and species that follow include features found in each taxon as a 
whole, including those populations and species occurring outside Ecuador 
and Peru. Phylogenetic studies applicable to species in this treatment may 
be found in Pipoly (1987) and in the forthcoming Flora Neotropica treatment. 


Cybianthus Mart., Nov. Gen. Sp. PI. 3:87. 1831. nam. ettypnsc(ms.\G. Agoscini, 
Acta Biol. Venez. 10:l4l. 1980.; Pipoly, Mem. New York Bot. Gard. 43:46. 1987; 
Pipoly, Ann. iVIissouri Boc. Gard. 79:9 13.1 992. Type specii-s: C. penduUflorus Martius. 

Terrestrial or epiphytic, monoaxial or polyaxial, dioecious, monoecious 
or polygamous shrubs or trees to 1 5 m tall. Roots positively geotropic or dia- 
geotropic. Branchlets glabrous, glandular-granulose, dendroid- and stellate- 
tomentose, furfuraceous- or ferrugineous-stipitate-lepidote. Leaves sessile or 
petiolate, alternate, subopposite, or pseudoverticillate, the venation 
camptodromous or rarely acrodromous; petioles obsolete or when present, 
canaliculate or marginate, tapering gradually to the base, or abruptly swol- 
len toward the base, here termed "pulvinate." Inflorescence: staminate, pistil- 
late, bisexual or polygamous, lateral (axillary), a simple raceme, panicle of 
racemose or spicate (rarely corymbose) branches, a pleiochasium, or an in- 
determinate umbel appearing racemose. Flowers functionally unisexual or 
bisexual, 3-6(-7)-merous; calyx cotyliform to cupuliform, the lobes im- 
bricate, valvate or aberrantly contorted, basally connate 1/5-2/3 their length, 
abaxially glabrous, glandular-granulose, ferrugineous stipitate-lepidote, or 
translucent-lepidote, adaxially glabrous, epunctate or prominently orange, 
red or black punctate, the lobes entire to erose-fimbriate, glabrous or glan- 
dular-ciliate; corolla rotate, subrotate, cupuliform or campanulate, rarely 
infundibuliform or salverform, the lobes imbricate or valvate, basally con- 
nate 1/5-3/4 their length, abaxially glabrous, glandular-granulose, or fer- 
rugineous stipitate-lepidote, adaxially glandular-granulose at least at the 
junction of the tube and lobe, the margin entire to erose-denticulate, gla- 
brous, glandular-granulose or rufous glandular-papillate; stamens and staminodes 
adnate to corolla tube at least 2/3 their length, the filaments variously con- 
nate to form a tube, the staminal tube adnate to the corolla tube or at times 

22 SiDA 18(1) 

developmentally fused with it (thus the stamens appearing epipetalous), 
bearing fleshy lobes alternate with the apically free portions of the filaments 
or not, the anthers erect or distally curved, ovate, widely ovate, or triangu- 
lar, basifixed or dorsifixed, apically acute, rounded, truncate, emarginate or 
minutely apiculate, the apiculum erect, proximally or distally curved, ba- 
sally truncate, cordate, or rarely hastate, deshiscent by apicaly pores, confluent 
apical pores (birimose), or by wide or narrow longitudinal slits; pollen tricolporate, 
psilate; staminodes morophologically similar to the stamens but greatly reduced 
in size, the antherodes at times producing abortive pollen; pistl obnapiform, 
ellipsoid, umbonate or obturbinate, the ovary sparsely to densely translu- 
cent glandular-lepidote, the style glabrous, the stigma capitate, capitate- 
lobate, or punctiform, persistent or early caducous, the placenta free-cen- 
tral, carnose, umbonate or globose, the ovules campyiotropous ( l-)2-5(-7), 
uni- or biseriate; pistillode conic, lageniform, obturbinate or irregularly shaped 
vestigial pistillode, the pistillode hollow or bearing a sterile placenta, rarely 
absent. Fruit drupaceous, 1(— 2)-seeded, the endosperm translucent, non- 
starchy, the embryo small, linear, flexuous, erect or curved, longitudinal or 
transverse, the cotyledons not well-developed. 

Distribution. — One hundred sixty-seven species; Nicaragua, Costa Rica 
south through Panama to the Andes southward to Bolivia, from Colombia 
eastward across Venezuela and Brazil and the Guianas, then southeastward 
to the Atlantic coastal forests of SE Brazil. 

Ecology. — Members o'i Cybianthus '<iK principally riparian, occurring only 
in primary forests or rarely in somewhat disturbed ones, and thus, may serve 
as indicators of environmental quality. Throughout the range of the genus, 
its members are known from wet tepuf savannas, moist scrub, cloud and 
elfin forests (including "ceja de selva"), subparamo thickets, montane, premontane, 
pluvial, wet and moist forests, paramo, jalca, igapo, varzea, varillal, campinas, 
campo rupestre, restinga, cerrado, and caatinga vegetation types. In Ecua- 
dor and Peru, the majority of the species occur in lowland and premontane 
forests at the junction of Hylaea and the eastern slopes of the Andean Cor- 
dillera (see ECOLOGY section). 

Cybianthus is most closely related to the paleotropical lianous genus Embelia 
(Pipoly 1 987), and cladistically defined by the unique glandular-granules at 
the corolla lobe and tube junction. For practical purposes of identification, the 
combination of lateral racemes or spikes, or racemose or spicate panicles, and 
filaments which are shorter than the corolla, connate at least 1/4 their length, 
and adnate to the corolla at least 1/3 its length, allows for easy recognition. 


1. Corolla corylitorm, cupLiliform, campanulate, or rarely salverForm; anthers 
longer than wide, disrally recurved, apically acure or minnrely apiculate. 

PiPOLY, Cybianthus in Ecuador and Peru 23 

2. Branchlets glabrous or glandular-granulose; anthers dehiscent by narrow 

longitudinal slits I. Microconomorpha 

2. Branchlets ferrugineous romentose or stipitate-lepidote; anthers dehis- 
cent by wide longitudinal slits. 
3. Branchlets and calyx ferrugineous stipitate-lepidote; abaxial corolla 

surface glabrous near margin II. Conomorpha 

3. Branchlets ferrugineous romentose, calyx glandular-granulose or gla- 
brous, rarely ferrugineous romentose; abaxial corolla surface glandu- 
lar-granulose near margm III. Laxiflorus 

1 . Corolla rotate to subrotate; anthers wider than long, erect, apically rounded 
to truncareor emarginate. 

4. Petioles abruptly swollen basally; anthers dorsifixed, longitudinally 
5. Plants monoaxial; anthers erect, not versatile. 

6. Stem glandular papillate, at times with hydropotes, bearing cataphylls 
apically and at times, alternating with the leaves; leaves without 
subepidermal fibers; flowers 4- or 5 -merous; corolla lobes epunctate 
or inconspicuously pellucid or orange punctate, glandular-granulose 
within; stigma large, capitare-lobate, early caducous, the lobe mar- 
gins fimbriate IV. Comomyrsine 

6. Stem with malpighiaceous hairs, without cataphylls, but at times 
with aborted leaves (pseudocataphylls); leaves with numerous paallel 
subepidermal fibers (most easily seen adaxially); flowers 3-merous; 
corolla lobes prominently black punctate, maculate, glabrous within 
except at lobe and tube junction; stigma small, capitate-lobate, per- 
sistent, the lobe margins entire V. Triadophora 

5. Plants polyaxial; anthers versatile VI. Weigeltia 

4. Petioles obsolete or not abruptly swollen basally; anthers basifixed, poricidally 
7. Leaves sessile, apically mucronate, basally auriculate, the margins 

scarious; staminal tube merely adnate to corolla tube VII. Grammadenia 

7. Leaves petiolate, apically acute, acuminate or caudate, basally acute, 
attenuate or cuneate, the margins opaque; staminal tube development- 
ally fused to corolla tube, the stamens thus appearing epipetalous 
VIII. Cybianthus 

I. Cybianthus subgenus Microconomorpha (Mez) G. Agostini, Acta Biol. 
Venez. 10:150. 1980; Pipoly, Wrightia 7:235. 1983. Conomorpha A. DC. subgenus 
Microconomorpha Mez inEngl., Pllanzenr. IV. 236(Heft 9):251. 1902. Microconomorpha 
(Mez) Lundell, Wrightia 5:349. 1977. Type Spi'cii^s: Conomorpha verticillata Zahlbr., 
Ann. K.K. Naturhist. Hofmus. 7:3. 1892, non Mez (1902). = Cybianthus pastensis 
(Mez) G. Agostini (lectot'i-pi,: Agostini, Acta f^iol. Venez. 10:150. 1980). 

Cybianthus Mart, subgenus Iteoides G. Agostini, syn. nov.. Acta Biol. Venez. 10:148. 
1980. Type Species: BadiiLi itcoicks Benth., PI. Hartw. 217. 1896. Conomorpha iteoides 
(Benth.) Mez in Engl., Pflanzenr. IV. 236(Hef't 9): 254: 1 902. Type Spiiciiis: Cybianthus 
iteoides (Benth.) G. Agostini, Acta Biol. Venez. 10:149. 1980. 

Terrestrial monoecious, clioecous or polygamous j7:??7/^i or sttiall trees. Bark 
smooth to slightly fissured, light brown, thin. Root positively geotropic. 
Trunk distinguishable, leptocaulous, the growtli dynamics following Rauh's 

24 SiDA 18(1) 

Architectural Model (Halle et al. 1978). Branchlets thin, terete, densely ter- 
rugineous glandular-granulose, the granules often stipitate. Cataphylls and 
pseudocataphylh absent. Leaves pseudoverticillate; blades petiolate, often with 
translucent glandular lepidote scales. l}ifloresce)ice a simple raceme or bipin- 
nate panicle, staminate, pistillate, or polygamous, the peduncle 1—4 cm long, 
densely glandular-granulose; mflorescence and floral bracts, perianth and 
pistil bearing prominently raised red or black punctations; infforescence bracts 
large, often foliaceous and persistent; fioral bracts linear-lanceolate, the margins 
glandular-ciliate, caducous, the pedicels erect, accrescent in fruit. Stami- 
nate, pistillate and bisexual flowers monomorphic (similar in shape), the staminate 
the largest, the pistillate the smallest in size, white to yellowish-green, (4—) 
5(— 6)-merous; calyx cotyliform, the lobes valvate, the margins densely glandular- 
ciliate; corolla cotyliform to campanulate, the lobes imbricate, glabrous without 
except glandular-granulose near the margin, glandular-granulose over the 
entire surface within; stamens and staminodes with a conspicuous staminal 
tube, the apically free portions one to three times longer than the anthers, 
the anthers elongate-triangular to ovate, prominently curved distally, apically 
obtuse to apiculate, basally cordate to hastate, clorsifixed 1/3 to more than 
1/2 length from base, dehiscent by narrow longitudinal slits, the staminodes 
producing abortive pollen grains; pistil in pistillate and bisexual flowers 
obturbinate, the ovary densely translucent-lepidote, the style thick, trun- 
cate, the style punctiform, the placenta umbonate, bearing 3—4 uniseriate 
ovules immersed in placental tissue, but exposed apically by placental pores; 
pistillode similar to pistil but reduced in size, hollow or bearing 2 abortive 
ovules. Fn/it drupaceous, 1 -seeded, the exocarp thin, prominently black punctate. 
Subgenus Mtcroconomorpha contains 5 species, of which one, Cybianthus 
pastensis (Mez) G. Agostini, is known from Ecuador and Peru. I earlier indi- 
cated that there was no evidence to support subgenus Iteoides as a separate 
entity (Pipoly 1987), so it is treated here in synonymy under subgenus 
M icroconomorpha. 

1. Cybianthus pastensis (Mez) G. Agostini (Fig. 3c), Acta Biol. Venez. 
10:151. 1 980. Conomorpha pastensis Me/ in Engl., Pflanzenr. IV. 236(Hefc 9):252. 
1902. Al/rmw/mor/;A<^/wmTO/.v(Mez)LLinclell,Wrigluia 5:349. 1977. Type: COLOMBIA. 
Narino: "Paramo de Purugai, Prov. cie Pasco," 2,500 m, 1866 (stam. fl),_/. Triana 
2585 (i.F.CTOTypK (Pipoly 1983a): W; isc)lectotypi;s: C, COL, G, P). 

Aiyrsine verticillcita C Prcsl, Reliq. Haenk. 2:64. 1835. Conomoyphci vtrthillata {C. Presl) 
Mez in Engl., Pflanzenr. IV. 236(Hcfc 9):252. 1902, mm Zahlbr. (1892). Conomorpha 
prest//).[\ Macbr., Cantlollea 5:398. I 934. Mkrnconomoipljci verticillcita (C. Presl) Lundell, 
Wrighria 5:349. 1977. Tyhi;: PERU. HcAnuco: without further locality, without date, 
(stam. fl.), T. Haerike 98 (i.hctotyph (Pipoly 1983a): PR; isolectotypes: HAL, W). Non 
Cybianth//s verticillatns (Veil.) G. Agostini, Acta Biol. Venez. 1():16S. 1980. 

Coiiomorpija verticillata Zahlbr., Ann. K. K. Naturhist. Hofmus. 7:3. 1892, non C. Presl 

PiPOLY, Cybianthus in Ecuador and Peru 25 

(1835) Cmwnwrpha jehkii Mez in Engl., Pflanzenr. IV. 236(Heft 9):25l. 1902, nom. 
superfl. Corioinorpha preslii ].V. Macbr. \-c\t. jelskii (Mez) J.F. Macbr., Field Mus. Nat. 
Hist., Bot. Ser. 13:201. 1959- Muroconomorpha jehkii {Mfi) Lundell, Wrightia 5:349. 
1977. Type: PERU. CajaxMARCa: Cutervo, Apr. 1879 (stam. fl.), C vonjelski 11 (ho- 
lotypk: W; piioto and fragment, F, F Neg. 31980). 

Conoiiwrpha (jeiitalei Mez in Engl., Pflanzenr. IV. 236(Heft 9):252. 1902. Microcono?Horpha 
dentata (Mez) Lundell, Wriginia 5:3-49. 1977. Typi:: ECUADOR. Pichincha: In cor- 
dillera from Quito to Tungurahua, 2, 000-3, 000 m, 1857-9 (stam. & bisex. fl.), R. 
Spruce 3 173 (lpctotype (Pipoly 1983a): K; lsolec:t()TYPF,s: BM, BP, C, CGE, GH, GOET, 
LD, LE, F Neg. 22956). 

Conoiiwrpha q/ieni folia Mez in Engl., Pllanzenr. IV. 236(FIeft 9):253. 1902. Microamnniorpha 
quercifoiia (Mez) Lundell, Wrightia 5:349. 1 977. Type: PERU: without locality, without 
date (stam. \\.),J. Pavon s.n. (lectotype (Pipoly 1983a): G). 

Conomorpha panamensis Lundell, Wrightia 5:290. 1976. Microconomorpha panamensis (Lundell) 
Lundell, Wrightia 5:349. 1977. Cyhianthus morn G. Agostini, Acta Biol. Venez. 10: 154. 
1980. Type: PANAMA. Chiriqui: Cerro Pando, on continental divide and Panama- 
Costa Rica border, ca. 16 km W of Hato del Volcan, 2,000-2,482 m, 20 Jul 1975 
(stam. fl.), S. Mon & A. Bnlten 7292 (hoeotype: LL-TEX; Isotype: MO). 

Shrub or tree to 6 m tall. Branchlets and inflorescence densely ferrugineous 
glandular-papillose, the branchlets angulate to prominently ridged, (1.5-) 
2-3 mm diam. Leaves pseudoverticillate; blades membranaceous to coria- 
ceous, narrowly oblanceolate to oblong or obovate, (3.0— )6. 5—14. 5(— 21.0) 
cm long, (1.2-)2-4.5(-6.8) cm wide, apically attenutate, acute or acumi- 
nate, basally cuneate, not decurrent on the petiole, prominently punctate 
and minutely ferrugineous stipitate-papillose above and below, the midrib 
impressed above, raised below, the secondary veins 7-15 pairs, prominently 
raised below, the margin undulate, lobate, crenate or dentate, rarely subentire; 
petioles marginate, (0.2-)0.5-2(-2.7) cm long, densely glandular-papillose. 
Staminate, pistillate or polygamous inflorescence: monomorphic, erect or lax, a 
simple raceme, 1.8-5.5 cm long, the rachis thin to thick, densely glandu- 
lar-papillose; inflorescence bracts chartaceous, obovate to elliptic, (4.3— )6- 
11 mm long, 3-7 mm wide, apically acute to acuminate, basally cuneate, 
densely and prominently red punctate; floral bracts chartaceous, linear-lan- 
ceolate, (0.8-)1.4-2.2(-7) mm long, 0.6-0.8 mm wide, apically attenuate, 
caducous; pedicels cylindrical, (1 .3-)2-7(-7.5) mm long, densely glandu- 
lar-papillose. Floivers (4-)5-merous, white to yellowish-green; calj^ chartaceous, 
shallowly cotyliform, (0.6-)0. 8-1.1 (-1.5) mm long, unequally divided, the 
tube 0.2-0.5 mm long, the lobes suborbicular to very widely ovate, (0.4-) 
0.6-1 mm long and wide, rounded to acute apically, glabrous, densely and 
very prominently orange or black punctate, the margin subentire to erose- 
dentate, densely glandular-ciliate; corolla chartaceous, cotyliform, 2-2. 6(- 
3.6) mm long, the tube 0.2-0.3 mm long, the lobes ovate to narrowly ovate, 
1.7-2.3(-2.8) mm long, 0.8-1.3(-1.5) mm wide, highly reflexed at anthesis, 
apically rounded to obtuse, prominently orange or black punctate; stamens 

26 SiDA 18(1) 

and staminodes 1-1.6(— 2.5) mm long, the staminal and staminodial tube 
0.7—1.8 mm long, the apically free portions of the filaments 0.3—0.7 mm 
long, the anthers elongate-triangular, 0.6-1.2 mm long, apically obtuse, 
basally hastate, the connective red punctate ventrally and dorsally, dorsifixed 
ca. 1/3 to slightly less than 1/2 from base; pistil and pistillode 1.2—1.8 mm 
long, the ovary (0.6-)0.8— 1 mm long, 1—1.3 mm diam., densely translu- 
cent glandular-lepidote, the style thick, 0.5-0.8 mm long, the stigma punctiform, 
the pistillode hollow or containmg one abortive ovule. Fruit globose, green, 
then red, then black at maturity, 3—4 mm diam. when dried exocarp thin, 
prominently pellucid punctate. 

Distribution. — Costa Rica to Colombia, southward to Peru, from 1,500- 
3,200 m elevation. 

Ecology CI nd conservation status. — Cyhianthus pastensis is known from premontane 
and montane pluvial and cloud forests, and at elfin forest margins. Popula- 
tions in areas exposed to winds have more coriaceous leaves and shorter stature, 
frequently as small as one meter in height. The wetter the habitat, the more 
membranaceous the leaves become, and the longer the inflorescences. Fieldwork 
in Colombia has shown that populations may contain six individuals per 
hectare, and that the population rapidly dwindles in areas of disturbance. 
Owing to population dynamics thus far observed, Cybianthiis pastensis should 
be considered threatened. 

Etymology. — The specific epithet refers to the area from which the type speci- 
men was collected, near the city of Pasto, Department of Nariiio, Colombia. 

Representative specimens examined. COLOMBIA. Antioquia: Mpio. Urrao, Parqiie 
National Natural "Las Orqui'deas," Vereda Calles, Permanent Premontane Rainforest In- 
ventory Plot, right bank of Rio Calles, 06" 32' N, 76° 19' W, 1 ,450-1 ,=)()() m, 29 Nov 
1993 (fr),./. Pipoly. A. Cogolloei al. 17322 (BRIT, COL, JAUM, MO), limits of Parqtie Las 
Orqui'deas, left bank of Ri'o Calles, 1 ,450- 1 ,500 m, 30 Nov 1 993 (ster.),/. P/po/y. A. Coi^iillo 
et al. 17376 (BRIT, JAUM, MO); near limit of Parque Las Orqui'deas, Alto de Palmitas, 
ca. 1 km from INDLRENA C;abana Calles, 1,300-1,400 m, I Dec 1993 (ster.),^. P//?()/y. 
A. Cogollodcd. I75()X n323 (BRIT,JAUM, MO), 2 Dec 1993 (fl bud),./. P/poIy. A. Cogdlo 
etal. 17334 (BRIT, COL, JAUM, MO), Right bank of Rio Calles, 1,350-1,450 m, 7 Dec 
1993 (stam. fl), /. Pipoly et al. 17881 (BRIT, COL, JALJM, MO); Along trail to Finca La 
Quince, above LJrrao, 06° 30' N, 76° 10' W, 2,500-2,800 m, 21 Nov 1988 (stam. H), G. 
McPhersoii cY al. 13212 (BRIT, HUA, MO); Mpio. Frontino, Region de Murrf, ca. 1 3 km 
from Nutibara, 06" 40' N, 76" 20' W, 2,000 m, 9 Dec 1 988 (pist. fl, fr), G. McPher.m/ d al. 
13397 (BRIT, HLJA, MO). Norte de Santander: San Antonio, W of Cali, near summit of 
Cordillera Occidental, 1 ,9800- 2,350 m, 26 I'eb-2 Mar 1 939 (stam. fl), E. Klll/p&A. Gania 
33886 (A, S, US). ECUADOR. Azuay: Chiguinda, on E slopes of cordiUera E of Sigsig, 
03° 12' S, 78° 36' W, 1 ,600-1 ,800 m, 1889 (stam. fl), /■ Lehmarni 5 143 (K-2 sheets). Carchi: 
Paramo de Achupallas, 01° 46' S, 78° 33' W, 2, 000-3, 000 m, 1899 (stam. fl), F. Lehmaiin 
6202 (K-2 sheets); From Prima Vera about 6 hrs. hike up Ri'o Gualchan Drainage to Nilo 
Ortiz' shelter, 00" 50' N, 77° 72' W, 1 ,9_^0-2,200 m, 7-8 Jun 1993 (fl bud),/. Bradford et 
al. 53 (BRIT, MO, QC;NE). Loja; Cerro Bangala, ca. 10 km E of Yangana, 2,500-2,700 m, 
I80ct 1988(pist. ri,fr),(7. H./r//w(,'253 /.)' (GB), (stam. fl), G7 Harling 23334 {GB)\CAmu'm 

PiPOLY, Cybianthus in Ecuador and Peru 27 

Loja, Carretera Loja-Zamora, at high point, 03° 58' S, 79° 04' W, 2,400-2,600 m, 23 Dec 
1991 (pist. fl), D. Ri/hioetal. 2232 (BRIT, xMO, QCNE); Loja, 3,500 m, 1 Dec 1876 (stam. 
fl), E. Andre 433 1 (F, K, NY); Divide between Quebrada Jipiru and E fork of Rfo Zamora, 
W slope of Cordillera de Zamora (El Condor), 9 km E of Loja, 04° 00' S, 79° 06' W, 2,700 
m, 19 Feb 1945 (stam. fl), F.R. Fosherfi & M. Giler 2.3119 (NY, US); Loma de Loro, 6 km S 
of Saraguro, on Rd. to Loja, 3,200 m, 1 1 Feb 1985 (stam. fl), G. Harling & L. Andersson 
21594 (AAU, S); Saraguro-Loja Rd., km 1 2.4, turnoff toward Fierro Urco, Km 3.8-7.1, 
03° 42' 33" S, 79° 18' 03" W, 3,120-3,390 m, 7 Dec 1994 (pist. fl), P.Jorgenseii et al. 1297 
(BRIT, LOJA, QCA, QCNE); W slope of Nudo de SabaniUa, ca. 8 km above Yangana on 
Rd. to Valladolid, 2,300-2,500 m, 2 Apr 1985 (bisex. fl, fr), G. Harling & L. Andersson 
23540 (GB); Cerro Toledo, Rd. to La Torre, ca. 7 km SE of Yangana, 2,500 m, 7 Apr 1985 
(stam. fl), G. Harling & L. Andersson 23842 (GB). Morona-Santiago: Between Campanas 
and Arenillas, along Ri'o Tintas, 10 leagues SE of EI Pan, 2,195 m, 13 Jul 1943 (stam. fl), 
_/. Steyermark 33642 (F, NY); Above Mirador, 2,375 m, 9 Sep 1943 (stam. fl),J. Steyermark 
'53897 (F, NY). Napo: 10 km W of Cuyuja, along Quito-Lago Agrio Rd., 00° 25' S, 78° 
00' W, 2,700 m, 3 I Apr 1983 (stam. fl), H. Balslev4293 (AAU, QCA); Salcedo-Napo Rd., 
2,390-2,590 m, 7 Feb 1977 (fr),J, Brandbyge 42093 (AAU, QCA); E of Borja, Cerro Antisana, 
28 Jul I960 (stam. fl), P. Grubb et ul. H)7 3 (OXF, NY); Sta. Barbara Scumbios, 00" 22' S, 
77° 10' W, 2,700 m, 10-15 Feb 1959 (stam. fl), /.. Holm-Nielsen 681 8 {AAU ,QCh);^-l2 
km ESE of Sta. Barbara, 00° 40' N, 77° 30' W, 2,780-2,880 m, 1 1 Jan 1985 (stam. fl),/, 
Luteyn & E. Cotton llOll (GB, QCA, NY, VEN); Paso de Guamani, Rio Chalpi, at bridge 
on Papallacta-Baeza Rd.., 2,800 m, 6 May 1967 (stam. fl), B. Sparre 13940 (GB); Canton 
Quijos, Sierra Azul (Agri'cola Industrial Rfo Aragon), 00° 40' S, 77° 55' W, 2,300 m, 2 
May 1992 (fr), A. Alvarez et al. 381 (BRIT, MO, QCNE), (fr), A. Alvarez et al. 412 (BRIT, 
MO, QCNE), Campamento Estero Chico, 00° 41' S, 77° 56' W, 2,500 m, 18 Jun 1992 
(fr), A. Alvarez et al. 490 (BRIT, MO, QCNE); Sierra Azul, Cordillera de Huacamayos, 00° 
41' S, 77° 54' W, 2,500-2,700 m. 10 Feb 1994 (fr), A. Alvarez et al.l330 (BRIT, MO, 
QCNE). Tungurahua: On Patate-Triunfo Rd., 01° 18' S, 78° 25' W, 2,950 m, 5 Nov 
1983 (stam. fl),./, Brandbyge & A. Barford 42306 (AAU, QCA, QNA, S). Zamora-Chinchipe: 
Rd. from Loja "to Zamora, km 14, 00° 04' S, 79° 09' W, 2,750-2,770 m, 19-20 Apr 1973 
(stam. fl), L. Holm-Nielsen et al. 3965 (AAU, QCA). PERU. Amazonas: Prov. Luya, 
Camporredondo-TuUanya, trail to Cerro Huicsocunga, 2,350 m, 3 Sep. 1989 (A), C Diaz 
&J. Campos 37 11 (MO, USM); Parte aita de las Montaiias de Galeras, 2,000-2,500 m, 20 
Jun 1991 (fl bud), C. Diaz et al. 4448 (BRIT, MO, USM). Ayacucho: Prov. La Mar, E 
massif of Cordillera Central, opposing the Cordillera Vilcabamba between Tambo San Miguel, 
Ayna and Hacienda Ltusiana, 12° 45' S, 73° 53' W, ca. 30 km SW of Hacienda and Rio 
Apurimac, 21 Aug 1968 (bisex. fl), T. Dudley 119 15 (F, NA, US). Cajamarca: Prov. Cutervo, 
10 km NW of Socota, 3,200 m, 10 Dec 1938 (stam. fl.), H. Stork & X. Horton 10134 (F); 
San Andres de Cutervo, Parque Nacional de Cutervo, "Jalca," trail to Laguna "EI Pileo," 
2,680 m, 15 Mar 1989 (fl, fr), C. Diaz et al. 3330 (AMAZ, MO, USM); Prov. Jaen, E side 
of Cordillera E of Huancabamba, 2,400-2,600 m, Apr 1942 (fr), A. Weherbai/er 6099 (F, 
GH, US). Huanuco: Prov. Pachitea, region of Pucallpa, W part of Sira Mountains and 
adjacent lowland, ca. 26-28 km ESE from Puerto Inca, 09° 25' S, 74° 43' W, 2,210 m, 15 
Aug 1988 (stam. fl), B. Wallnbfer 1 1-16888 (BRIT, MO, W, WU, USM); SW slope of Rfo 
LluUa Pichfs Watershed, on the ascent of Cerros del Sira, top of first cumbre between camp 
4 (Peligroso) and camp 5 (Tabano), 1,680 m, 31 Jul 1969 (ster.), T. Dudley 13513 (NA). 
Lima: Prov. Lima, Lima, without date (fr),y, Pavon s.n. (K). Madre de Dies: Prov. Manii, 
Cerro dePanciacolla, RfoPlotoa, 10-15 km NNWof Shintuya, 12°35' S, 71°18' W, 1,000- 
1,400 m, 15 Dec. 1985 (fr), R. Poster et al. 10860 (F, MO, USM). 

Cybianthus pastens is may be easily recognized by its pseudoverticillate leaves 

28 SiDA 18(1) 

with variously serrate or incised margins, the very fine inflorescence rachis 
and minute flowers, and tlie stipitate papillae of the branchlets, leaves, petioles 
and inflorescence rachises. The prominent ridges of older branchlets and 
the swollen pseudoverticels of leaf scars are also distinctive. 

II. Cybianthus subgenus Conomorpha (A. DC.) G. Agostini, Acta Biol. 
Venez. 10:150. 1980; Pipoly, Ann. Missouri Bot. Gard. 79:908-957. 
1 992. CoNoi/iorphj A. DC, Trans. Linn. Soc. Loncli)n, Bor., 17:1 02. 1 83-1; Co)!(ini(>rphii 
sect. Eiimnniwiylihct Mit|., Stirp. Surinam. Select. 111. 1850; Couoiiiorpha subgenus 
Eucommorpha Mcz in Engl., Pflanzenr. IV. 236(Heft 9):254. 1902. Typi- Sppcirs: Cononnnphc) 
ohlongifolia A. DC. = Cybianthus ohlovgijoliiis (A. DC.) G. Agostini (lkcioiype: by 
Agostini, Acta Biol. Venez. 10:1 VI. 1980). 

CoiiDDioyphii sect. Aai)ifiinorpl:ki Miq. in Mart., I'l. Bras. 1 0:304. 1856. Type Spi;(:ii;s: Conoinoypbu 
heteranthd Benth. (lec;t()[YPe: Agostini, Acta Biol. Venez. 10:151. 1 980)= Cybhnithus 
y^/iyam-Hsis (A. DC^. ) Mic]. subsp. i^//yai/ti/s/s. 

Terrestrial dioecious, bisexual, polygamous, or rarely, monoecius shrubs 
or trees. Roots positively geotropic. Bark smooth or fissured, brown, or beige, 
rarely with significant amounts of cork. Ih/nk distinguishable, leptocaulous, 
the growth dynamics following Rauh's or rarely, Aubreville's Architectural 
Model (Halle et al. 1978). Branchlets thin to moderately thick, terete or ridged, 
densely to moderately covered with ferrtigineous stipitate-lepidote scales, 
the scales at times appressed, rarely glabrescent. Cataphylls and pseudocataphylls 
absent. Leaves alternate, rarely approaching pseudoverticillate (C peruvianus), 
petiolate, covered with ferrugineous stipitate lepidote scales, often glabrescent 
above; petioles canaliculate, marginate, or rarely winged. Inflorescence race- 
mose, spicate or paniculate, the panicles with racemose branches, rarely a 
solitary flower; inflorescence bract small, lanceolate, early caducous; rachis 
erect or lax, straight or rarely tortuous, ferrugineous stipitate-lepidote; floral 
bracts deltate, lanceolate or ovate, ferrugineous stipitate-lepidote, inserted 
at the base of the pedicel; pedicels cylindrical, at times clavate in frtiit or 
absent, erect, apically recurved, pendent, or nodding, at times accrescent 
in fruit. Flowers unisexual, rarely bisexual, dimorphic, (3-)4-5(-6)-merous; 
calyx cotyliform, cupuliform, crateriform, urceolate or patelliform, the lobes 
valvate, epunctate or with prominent (raised and blisterlike), conspicuous 
(readily visible buy flat), or inconspicuous brown, red or black punctations, 
the margin entire, rarely crenulate or erose; corolla campanulate to cupuliform, 
rarely salverform or tubiform, the lobes erect or reflexed, rarely cucuUate, 
valvate or imbricate, ferrugineous stipitate-lepidote or glabrous and epunctate 
or prominently, conspicuously or inconspicuously pellucid, brown, or black 
punctate without, at times with a narrow line of glandular-granules along 
the margin, gland ular-granulose within, the margins enrire or rarely crenulate, 
glabrous or rarely glandular-granulose; staminodes resembling stamens but 
reduced in size, the tube conspicuous or inconspicuous, adnatc to the co- 

PiPOLY, Cybianthus in Ecuador and Peru 29 

rolla, lobate or elobate, the anthers ovate or triangular-ovate, rarely linear- 
lanceolate, rarely deltate, usually recurved distally, rarely erect, apically acute, 
or apiculate, rarely rounded, the apiculum dorsally, rarely proximally re- 
curved or erect, the base cordate, dorsifixed from near base to subversatile, 
the connective punctate or not; pistillode conic to lageniform, rarely ab- 
sent, translucent-lepidote or glabrous, hollow; pistil obnapiform, rarely conic, 
the ovary globose, lobed or with an apical apophysis, the style short, the 
stigma capitate-lobate, 2-3-lobed or punctiform; placenta cupuliform or 
cotyliform, the ovules 2-4. Fri/it subglobose, one(-two)-seeded. 

Cybianthus subgenus Conomorpha contains 44 species, 10 of which have 
been recorded from Ecuador and Peru. 


1. Branchlets with erect stipitate ferrugineous lepidote scales, their margins 
not appressed; leaf blades subbullate to bullate, the secondary veins some- 
what to deeply impressed above, prominently raised below; corolla inlundibu- 
liform or tubiiorm, or appearing so in bud. 

2. Branchlets flexuous, 4-5 mm diam.; leaf blades perpuncticulose above, 
sparsely lepidote below, the secondary veins 22-26 pairs, the margin ir- 
regular; inflorescence tortuous, pinnarely to bipinnately paniculate; co- 
rolla lobes prominently keeled, rugose without; anthers ventrally 

recurved 2. C. gigantophyllus 

2. Branchlets straight, 2-3 mm diam.; leaf blades not perpuncticulose above, 
ensely lepidote below, the secondary veins 8-19 pairs, the margin regu- 
lar; inflorescence erect, a simple raceme or poorly formed panicle consist- 
ing of basally clustered racemes; corolla lobes flat, smooth or verrruculose 
without; anthers dorsally recurved. 

3- Secondary veins 12-16; staminal tube epunctate; pedicels cylindric; 
calyx cotyliform; corolla membranaceous, infundibuliform, verruculose 
without; fruit smooth, 3.5-4.5 mm diam.; plants of premontane plu- 
vial forests, subparamo thickets and upper pluvial cloud forests, 1,000 

1,960 m elevation 3. C. occigranatensis 

3. Secondary veins 8-12; staminal tube punctate; pedicels obconic; calyx 
urceolate; corolla carnose, tubiform, smooth without; fruit costate, 7- 
15 mm diam.; plants of white sands or on sandstone, 150-180 (-1,500) 

m elevation 4. C. spichigeri 

1 . Branchlets with appressed ferrugineous lepidote scales, the margins appressed; 
leaf blades not subbullate or bullate, the secondary veins planar or slightly 
raised above, barely discernible or slightly raised below; corolla campanu- 
late to cupuliform. 

4. Leaf blades coriaceous, rarely chartaceous, the margins subrevolute to 

5. Leaf blades densely and prominently pustulate at maturity above, the 
secondary veins 24-28 pairs, inconspicuous below; inflorescence spi- 
cate (2-)6-l6 cm long; flowers subsessile, the pedicels 0.2-0.6 mm 

long; calyx deeply cupuliform 5. C. lepidotus 

5. Leaf blades essentially smooth or sparsely pusticulate at maturity above, 

30 Si DA 18(1) 

thf secondary veins 14—25 pairs, prominently raised below; inllorescence 

a raceme or jianicle witli 2-4 racemes branching from base, (2-)3-7 (- 

8) cm long; Bowers pedicellate, the pedicels ((>.')-)(). 9- 1 .5 mm lon^; 

calyx cocylitorm. 

6. Branchlets stibterete, 2-3 mm diam; leaf blades 1 .2-2(-2.5) cm wide, 
smooth above; petioles '3~7(-l()) mm long; staminate calyx carnose, 
1 .2—1.8 mm long; staminate corolla carnose, densely lepidote with- 
out, the scales overlapping, 3.2-3.'1 mm long, the tube equal to the 
staminal cube, the lobes symmetric; anthers ovate; fruit with fleshy 
exocarp; plants of montane and clotid forests on sandstone 6. C. laetus 

6. Branchlets terete, 3~4 mm diam.; leaf blades (2.6-)3.5-3 cm wide, 
pusticulate above; jietioles 10-15 mm long; staminate calyx 
chartaceoLis, 0.8-1.2 mm long; staminate corolla chartaceous, 
glabrous or sparsely lepidote without, the scales not overlapping, 
2.0-2.6 mm long, the tube shortet than the stammal tube, 
the lobes asymmetric; anthers linear-lanceolate; Iruic with thin 
exocarp; plants of lowland and lower montane forests on white 

sands 7. C. peruvianus 

4. Leaf blades chartaceous to membranaceous, the margins flat. 

7. Leaf blades membranaceous to subchartaceous; petioles 5-1 0(-l 2) mm 

long; inflorescence a simple raceme or rarely 2-branched at base, 1-3 

cm long; corolla salverform or campanulate; fruit globose. 

8. Branchlets angulate, 1 .5-2 mm diam.; corolla salverform, the stami- 
nate 2,2-2.6 mm; plants of premontane forests on sandstone and 
limestone, (244-)400-l ,200 m elevation 8. C. comperuvianus 

8. Branchlets terete, 2-] mm diam.; corolla campanulate, the stami- 
nate, 2.8-S.2 mm; plants of lowland igapo forests, 90-240(-700) 

m elevation 9- C. guyanensis stibsp. pseudoicacoreus 

7. Leaf blades chartaceous; petioles ( 1 0-) 1 3- I 7(-22) mm long; 
mflorescene a panicle with 2-8 racemes branched from base, 4-8 cm 
long; corolla chartaceous, infundibuliform or c(Jtyliform; fruit de- 

9. Branchlets, petioles, abaxial leaf blades, inflorescence antl calyx lobes 
moderately to densely lepidote, but the not scales overlapping; leaf 
blades smooth above at matutity; corolla cotyliform, the lobes ob- 
long to oblanceolate, flat, smooth without, conspicuously black 
l^tinctate, apically acLiminate; staminal and staminodial ttibe charta- 
ceous, conspicuous; anthers and antherodes obcordate, the apiculum 

distally recurved; pistillode conic; pistil lagenitorm 10. C. tinianae 

9. Branchlets, petioles, abaxial leaf blades, inflorescence and calyx lobes 
moderately to densely lepidote, the scales overlapping; leaf blades 
pustulate above at maturity; corolla infundibuliform, the lobes ovate, 
verruculose without, iiicons|Ticuotisly brown |uinctate, apically 
rounded; staminal and staminodial cube membranaceous, inconspicu- 
otis; anthers ovate, antherodes subdeltate, the apiculum proximally 
inllexed; pisiillode lageniform; |-iistil obnapiform I I . C. cuatrecasasii 

2. Cybianthus gigantophyllus Pipoly, (Fig. 2 A, 10). CandoUea 46:4 1 . 199 1 • 

Bafsapuertoand Moyobamba, 600-1,200 

PiPOLY, Cybianthus in Ecuador and Peru 


Fig. 10. CybicmthHi gigantophylliis Pipoly. A. Habit, siiowing flexuous branchlet, paniculate 
inflorescences. B. Pistillate flower, showing urceolate calyx and crenulate corolla lobe mar- 
gins. C. Pistdlate flower with one corolla lobe removed, showing cucullate corolla lobe 
apices, proximally recurved anrherodes, and capitate, lobed stigma. A— D, drawn from ho- 
lorype, by Peggy Duke. Figure reproduced from Pipoly, 1991- 

32 SioA 18(1) 

m, Aug-Sep 1933 (pist. tl), G. King 3165 (noi.OTYPE: US; isotypf.s; l\ G-2 sheets, 
GH-2 sheets, MO, NY, US). 

Tree to 4 m tail. Branchlets flexuous, prominently ribbed, 4-5 mm diam., 
moderately lepidote. Leaves alternate; blades chartaceous, elliptic, (15.5)17- 
27 cm long, (5.9— )7-9.1 cm wide, apically long-acuminate, the acumen 
1 .2-3.5(-4) cm long, pustulate, perpuncticulose and glabrous above, sparsely 
lepidote below, midrib slightly depressed above, prominently raised below, 
the secondary veins 22—26 pairs, slightly depressed above, prominently raised 
below, the margin irregular, flat, entire; petioles canaliculate, thick, (1.6—) 
2.7—4 cm long, ca. 3 mm diam., sparsely lepidote, prominently ridged below. 
Staminate inflorescence: unknown. Pistillate inflorescence: a pinnate to bipinnate 
panicle, 1.5-2.5 cm long, tortuous, the branches spicate, moderately lepi- 
dote; peduncle 0.3-0.5 cm long, floral bracts carnose, deltate, 0.8-0.9 mm 
long and wide, apically acute, margin crentdate basally, densely lepidote 
above and below; pedicels obsolete. Pistillate flowers 4-merous; calyx carnose, 
urceolate, 1.6—1.8 mm long, the tube 0.9—1 rnm long, the lobes widely 
triangular, 0.5—0.7 mm long, 1-1.2 mm wide, apically acuminate-apicu- 
late, the margin regular, entire, lepidote; corolla carnose, campantdate, 2.7— 
3.1 mm long, the tube 0.2—0.3 mm long, the lobes erect, 2.5-3 mm long, 
apically rounded to obtuse, prominently cucuUate, abaxially carinate, apically 
rugose and glandular-granulose along the margins without, inconspicuously 
black punctate, the margin glandular-gran tdose, erose-crenulate; staminodes 
2.3—2.5 mm long, the staminodial tube membranous, inconspicuous, 0.2— 
0.3 mm long, elobate, glabrous, the apical free portions of the filaments 
1.2—1.3 mm long, flat, the anthers ovate, 0.8-1 mm long, 0.6—0.8 mm wide, 
apiculate, the apiculum ventrally recurved, basally cordate, the connective 
epunctate; pistil obturbinate, 1.8—2 mm long, 1—1.3 mm diam., the ovary 
1.4—1.6 mm long, the stigma capitate, 3— 5-lobed, the placenta cupuliform, 
ovules 3, erect, the upper portions exposed. Fruit globose, 4—5 mm long 
and in diam., exocarp thin, black, inconspicuously pellticid punctate. 

Distribution. — Cybianthus gigantophyllus is known from the headwaters of 
the rios Marafion and Huallaga in San JMartin, and the Iquitos area, along 
the Ri'os Napo, Nanay and Amazonas in Loreto, at 130—500 m elevation. 

Ecology and conservation status. — Cybianthus gigantophyllus occurs in pri- 
mary terra flrme forests, and on white sands (varillal) of lowland Peruvian 
Amazonia. Given increasing pressure from deforestation, it should be con- 
sidered threatened. 

Etymology. — The specific epithet refers to the leaf size, one of the largest 
known for the subgenus. 

Local names. — Peru: "ukushnum," "wewe," "yakCisnum," "yakushnum" 

PiPOLY, Cybianthus in Ecuador and Peru 33 

Representarive specimens examined. PERU. Amazonas: Prov. Bagua, Dtto. Imaza, 
Comunidad Aguarana de Putuim (CAMPOU), anexo Yamayakat, Monte Alco de Putuim, 
450 m, 25 Aug 1994 (fl bud), C. D/az et al. 7007 (BRIT, HUT, MO, USM); Quebrada 
ChichijamEncsa, RioCenepa, 130 m, 7 Jun 1973(fr), £. Ancuash 580 {AMAZ, MO, NY); 
Vicinity Huampami, 5 km E of Valdivia, 04° 30' S, 73° 30' W, 200-500 m, 12 Aug 1978 
(fr), £. Ana/ash 1437 (AMAZ, BRIT, MO, NY, US). Loreto: Maquisapa, Upper Rio Nanay, 
Jul 1929 (fr), U. Willuims 1 182 (F); Prov. Maynas, Dtto. Sta. Man'a de Nanay, Caseri'a Mishana, 
halfway between Iquitos and Sea. Maria de Nanay, 03° 50' S, 73° 30' W, 130 m, 25 Feb 
1979 (ster.), A. Gentry &J. Aronson 25044 (AMAZ, MO); Dtto. Las Amazonas, Quebrada 
Yanamono, Explornapo Tourist Camp, above mouth of Ri'o Napo on Rio Amazonas, 9 Nov 
1979 (ster.), A. Gentry et al. 27952 (AMAZ, MO), 25 km NE of Iquitos, along Ri'o Amazonas, 
southern perimeter path, 1 10 m, 27 Sep 1990 (ster.),/ Fipolyetal. 72^97 (AMAZ, BRIT, 
MO, US, USM), 03° 20' S, 72° 55' W, 100-140 m", 15 Feb 1991 (ster.),/ Pipoly et al. 
13028 (AMAZ, BRIT, MO, USM), 03° 28' S, 72° 50' W, 106 m, 15 May 1989 (ster.), R. 
Vdsqiiez et al. 12 108 (AMAZ, MO, USM); Explornapo Tourist Camp, near Sucusari, along 
Ri'o Napo, 03°20' S, 72°55' W, 100-140 m, 22 Feb 1991 (ster.), J. Ptpoly et al. 13284 
(AMAZ, BRIT, MO, USM), 23 Feb 1991 (ster.),/ Ptpolyetal. 13423 (AMAZ, BRIT, MO), 
(ster.),/ Ptpolyetal. 13426 (AMAZ, BRIT, MO, USM), 1 Mar 1991 (ster.),/ Ptpolyetal. 
13931 (AMAZ, BRIT, MO, USM). 

Cybianthus gigantophyllus is most closely related to C. occigranatensh (Cuatrec.) 
G. Agostini and C. spichigeri Pipoly. However, the large, flat leaves, long 
petioles, and tortuous panicles allow for easy recognition. In the original 
description (Pipoly 1991), I described the pistillate corolla as tubular, when 
it is, in fact, campanulate. When the flower is in bud, the corolla appears 
tubular as it longitudinally extends above the calyx, then it gradually opens, 
with cucuUate apices. Within the tall terra firiiie forests on lateritic soils, it 
may be found above the flood line along small creekbeds. 

3. Cybianthus occigranatensis (Cuatrec.) G. Agostini, Acta Biol. Venez. 
10:155. 1 980. Conomorpha occigranatensis Cuatrec, Revista Acad. Colomb. Ci . Exact. 
8(31):320. 1951. Type: COLOMBIA. Valle Del Cauca: Cordillera Occidental, W 
slope, Ri'o Digua River Basin, left bank of Rfo San Juan, around Queremal region, 
small stream at km 51, 1,540-1,650 m, (stam. fl),/ C/iatrecasas 237 34 (iiolotypi;: 
F; isotype: COL). 

Shrub or small tree to 4 m tall. Branchlets straight, subterete, 2—3 mm diam., 
densely lepidote. Leaves alternate; blades membranaceous, elliptic to obo- 
vate, (4-)7.5-l4(-21) cm long, (2.5-)4-5(-7) cm wide, apically acuminate, 
the acumen 1 .2— 1 .5(— 3.0) cm long, basally acute, decurrent on the petiole, 
the midrib impressed above, prominently raised below, the secondary veins 
8—12 pairs, deeply impressed above, prominently raised below, the leaf strongly 
bullate, adaxial surface smooth, densely lepidote when young, becoming 
pusticulate and sparsely lepidote or glabrous with age, abaxial surface densely 
lepidote, but the scales not overlapping; petioles canaliculate, 1.0— 1.5(— 
1.8) cm long, densely lepidote, persistent. StaniDiate inflorescence a raceme or 
a panicle with 1-3 branches from the base, 4—8 cm long; peduncle, rachis, 

34 Si DA 18(1) 

branches and pedicels densely lepidote; peduncle 0.1-0.4 mm long; floral 
bracts membranceous, ovate, shorter than the pedicels, 0.7-1.1 mm long; 
0.4-0.5 mm wide, apically acute, densely lepidote abaxially, the margin 
entire; pedicels cylindric, thin, 1.5-6 mm long. Stamincite flowers 4-merous; 
calyx carnose, cotyliform, 0.8-1.0 mm long, the tube 0.2-0.3 mm long, 
the lobes triangular to deltate, 0.5-0.9 mm long, 0.4-0.8 mm wide, apically 
attenuate to an acute or round tip, sparsely lepidote without, glabrous within, 
conspicuously brown punctate, the margin lepidote; corolla membranaceous, 
campanulate, 2.4-2.7 mm long, the tube 0.7-0.8 mm long, the lobes ovate, 
1 .6—1 .8 mm long, 1 .0-1.3 mm wide, apically attenuate to a round tip, veraiculose 
without, smooth within, sparsely lepidote without toward iipex, apically 
glanduiar-granulose within and along margins, conspicuously brown punctate, 
the margins entire; stamens 1.8-1.9 mm long, the filaments 2.6-2.8 mm 
long, the tube membranaceous, inconspicuous, adnate to the corolla tube, 
elobate, the apically free portions 0.2-0.3 mm long, the anthers triangular, 
0.8-1.0 mm long, 0.5—0.6 mm wide, apically attenuate to an acute , dor- 
sally reflexed tip, basally cordate, dorsifixed just above base, the connective 
dark, prominently brown punctate; pistiUode lageniform, 1.3-1.5 mm long, 
densely translucent glandular-lepidote near the base. Bisexual and pistillate 
inflorescence: as in staminate but only rarely branched from base, 4—6 cm long; 
peduncle 0.1—0.3 cm long; floral bracts 0.5-0.8 mm long; pedicels 1.5— 
3.5 mm long. Bisexual and pistillate flowers as in staminate but calyx 0.8— 

1 .1 mm long, the tube 0.3—0.6 mm long, the lobes deltate to oblate, 0.4— 
0.6 mm long, 0.8—1 mm wide; the margin irregular, entire; corolla as in 
staminate but 2.4-2.6 mm long, the tube 1 .0-1 . 1 mm long, the lobes ovate, 
1.4—1.6 mm long, 0.8—1.1 mm wide. Bisexual flowers with stamens 1.6— 
1.8 mm long, the tube 1.0-1 .2 mm long, the apically free portions of fila- 
ments 0.2-0.3 mm long, the anthers 0.5—0.8 mm long, pistillode 1.5—1.8 
mm long. Pistillate flowers with staminodes 1 .6—1 .8 mm long, the tube 1 .0- 

1.2 mm long, the apically free portion of filaments 0.1-0.2 mm long, the 
antherodes 0.5-0.7 mm long; pistil obnapiform, 2.4-2.6 mm long, the ovary 
1.1 — 1 .2 mm long, 1. 1-1.2 mm diam., the style 1 .0 mm long, the stigma 
capitate, 2-lobed, to 0.2 mm long, the ovules 2-4, buried in the placenta 
below apical pores. Fr///> globose, 2.5-4 mm long, 3.5—4.5 mm diam., the 
endocarp smooth, the aril scanty and adnate to both seed and endocarp, the 
embryo straight, ca. 3 mm long. 

Distribution. — Panama (Darien), Colombia (Cordillera Occidental) and 
Ecuador (Esmeralda, Napo, Santiago-Zamora, Sucumbios), at 1 ,000— 1 ,960 
m elevation. 

Ecology aiul conservatu))! status. — Cyinaiitbus occigraiiatensis occurs in premontane 
pluvial forests, subparamo thickets and in upper pluvial cloud forests. Based 
on my observations of populations in subparamo thickets at the Antioquia/ 

PiPOLY, Cybianthus in Ecuador and Peru 35 

Choco interface in tiie Cordillera Occidental of Colombia, this species tol- 
erates disturbance well as long as the soil is not compacted. It is retricted 
to areas where rainfall exceeds 5,000 mm annually. At this time, the spe- 
cies does not seem to be threatened. 

Etymology. — The specific epithet refers to its principal range of distribu- 
tion, the Cordillera Occidental of Colombia and adjacent Ecuador. 

Specimens examined. PANAMA. Darien: S slojie of westermost summit of Cerro Tacaracuna, 
massif between Pucro base camp and Tacaracuna summit camj-i, 1,4()0— 1,600 m, 21 Jul 
1976 (stam. fl), A. Gentry et a l. 16867 (COL, LL-TEX, MO, PMA). COLOMBIA. Antioquia: 
Mcpio Frontino, km 13 Nutibara-La Blanquita Rd., Region de Mum', Alto de Cuevas, 
06° 44' N, 76° 23' W, 1 ,990 m, 6 Nov 19<S8 (fl bud),./. Zarucchi et al. 1201 (BRIT, HUA, 
MO); Mpio. Frontino, Vereda Venados, Parque Nacional Las Orquideas, sector Dos Bocas, 
confluence of Rio Venados and Rio Calles, ()6° 34' N, 76° 30' W, 29 Oc 1986 (stam. fl), R. 
Calkjas et al. 2737 (HUA, MO); M]iio. San Lui's, Autopisra Medelli'n-Santafe de Bogota, 
sector Rfo Samana, Rd. toward Vereda La Josefina, I (S Dec 1982 (stam. fl), A. Cogollo & C. 
Estrada 296 (COL, JAUM, MO); Mpio. Urrao, Parque Nacional Las Orqufdeas, Vereda 
Calles, Permanent Premontane Rainforest Inventory Plot, right bank of Rfo Calles, 06° 
32' N, 76° 19' W, 1,450 m, 26 Nov 1993 (ster.),^, Pipoly. A. Cogollo et al. 17 1 59 (BRIT, 
JAUM, MO), 27 Nov 1993 (ster.),^. Pi/mly, A. Cogollo et al. 17182 (BRIT, JAUM, MO), 
Range NW of Cabana de Calles, 1 ,450 m, 28 Nov 1993 (ster.), A. Cogollo etal. 7529 (BRIT, 
JAUM, MO), 1,450-1,500 m, 28 Nov 1993 (ster.),/ Pipoly etal. 17253 (BRIT, JAUM, 
MO), (ster.),/ Pipoly et al. 17281 (BRIT,JALJM, MO), 7 Dec 1993 (.ster.),/ Pipoly et al. 
17871 (BRIT, JAUM, MO), 9 Dec 1993 (fl. bud),/ Pipoly et al. 17979 (BRIT, JAUM, 
MO), Vereda Calles, Alto de Palmitas, ca. 1 km from Cabana de Calles, 1 ,700-1 ,750 m, 2 
Dec 1993 (ster.),/ Pipoly etal. 1 7542 (BRIT, JAUM, MO). Choco: Mpio. Itsmina, Quebrada 
Raspadura, between Raspadura and Quibdo, split of Rfo Atrato and Rfo San Juan drainage 
basins, ca. 05° 15' N, 76° 38'W, 18 Apr 1979 (fr), E. Forero & R. Jaramillo 5307 (COL, 
MO); Serranfa del Darien, along Colombian/Panamanian border, 1,400 m, 20 Jul 1976 
(stam. fl, bisex. fl), A. Gentry. H. Leon & L. Porero 16842 (COL, MO); without locality, 1866 
(fr),/ Triana 2589 (G). Huila: Rfo Suaza, SW of Alejandrfa, 1 ,670 m, 23 Aug 1 944 (stam. 
fl), E. Little 8532 (COL, US). Quindi'o: Mariquita, 1866 (stam. fl),^, Triami 2562 (P). 
Valle Del Cauca: Cordillera Central, 5 km N of Darien along Rd. toward La Guajira, Upper 
Rfo Calima, 03° 58' N, 76° 28' W, 1,550-1,700 m, 24 Jan 1986 (fl bud), B. Stein & L. 
McDacle 3284 (BRIT, HUA, MO); Finca Zungara, Corregimiento La Divisora, crest of Cordillera 
Occidental, W of Cali, 6 km N of Cali-Buenaventura Hwy, 03° 32' N, 76° 35' W, 1,960 
m, 12 Dec 1985 (ster.), A. Gentry etal. 53167 (COL, MO, US), 24 Mar 1986 (fr), A. Gentry 
etal. 53551 (COL, MO, US); Rfo Digua Drainage Basin, Piedra de Moler, 900-1,180 m, 
20 Oct 1943 (pist. fl, fr),/ Cuatrecasas 14918 (COL-2 sheets, F); Rfo Sanquininf, La La- 
guna, 1,250-1,400 m, 10 Dec 1943 (stam. fl),/ Cuatrecasas 15658 (COL, F, US); Monte 
La Guardia, La Carbonera Range, between Las Brisas and Alban, 1,950—2,000 m, 16 Ocr 
1946 (stam. fl),7. Cuatrecasas 2213 1 (COL, F, US, YEN); San Antonio, W of Cali, 1 ,900- 
2,350 m, 26 Feb 1 939 (stam. fl), E. KilUp&A. Garcia 33898 (A, BM, COL, F, NY, US); La 
Cumbre, 7 May 1 922 (stam. fl), P. Penmll5l47 (GH, K, NY, US). ECUADOR. Esmeraldas: 
San Lorenzo Canton, Reserva Etnica Awa, Parroquia Alto Tambo, Centro de la LJnion, Cation 
del Rfo Mira, 00° 52' N, 78° 26' W, 250 m, 22 Mar 1993 (fr), C. & M. Ai/lestia 1313 
(BRIT. MO, QCNE), Napo: Carrerera Nueva, Cotundo-Coca, 1 ,130 m, 5 Aug 1984 (pist. 
fl, fr), C. Doclson et al. 15115 (MO); Canton Archidona, 1 50 m NE of Caserio of Huamanf, 
right side of Carretera Hollfn-Loreto, 00° 43' S, 77° 36' W, 1,200 m, 9 Sep 1988 (fr), R 
Hurtado & D. Neill 235 (MO, QCNE), Cordillera de Guacamayos, Rd. to Archidona, Rfo 

36 SlDA 18(1) 

HoUin PeqLicno, primary forest on 90° slopes, 00° 38' S, 77" 48' W. 1 ,900 m, Aug I 990 
(stam. fl), W. Pcilacnis & E. fveire 4S99 (BRIT, MO, QCA); Canton El Chaco, Right margin 
of Rio Quijos, Finca "La Ave Brava," of Scgundo Pacheco, 00° 12' S, 77° 39' W, 1,800- 
1 ,900 m, 7- 1 Sep 1990 (fr), W PaLnws 5394 (BRIT, MO, QCNE); S slope of Volcan Reventador, 
left bank of Ri'o Reventador, between Rd. and trail to crater, 00° 07' S, 77° 30' W, 1 ,600- 
1 ,850 m, 1 1 Oct 1990 (stam. H), W. Paiacios 61 76 (BRIT, MO, QCNE), (stam. tl), W. Pa/aaos 
6187 (BRIT, MO, QCNE), (fr), W. Pa/aaos 6218 (BRIT, MO, QCNE); Proyecto Hidroelectnco 
(]oca, Punto ST4; right margm of Rfo Quijos, ca. 10 km S of Reventador, 00° 11' S, 77° 
39' W, 1 ,500 m, 3-5 Oct 1990 (pist. fl), W. PaU/os 58 H (BRIT, MO, QCNE), 08° 08' S, 
77° 30' W, 1,450 m, 6-10 Oct 1990 (pist. Il), W. Paiacios 6040 (BRIT, MO, QCNE); Yasum' 
National Park, Maxus Rd and pipeline construction project, km 15, 00° 3 1 ' S, 76° 32' W, 
250 m, 30 Jtin 1 994 (bud) N. P/tma/i 461 (BRIT, MO). Santiago-Zamora: Between Campanas 
and Arenillas, along Ri'o Tmtas, 10 leagues SE of El Pan, 2,195 m, 1 3 Jul 1913 (stam. fl), 
J. Steyer/fiark 53550 (NY). Sucumbios: Sendero toward Volcan el Reventador from km 
100 of Baeza-Eago Agrio Elwy, 1,900 m, 7 Oct 1990 (stam. ?i),J. Jaramillo & E. Grijalva 
12988 (QCA). 

Cybianth?/s occigranatensis is most closely related to C timanae Pipoly, but 
is easily distinguished by the fewer secondary veins of the coriaceous leaf 
blades, the campanulate corolla with verrucose, prominently black punc- 
tate lobes and attenuate apices, and obnapiform pistil. The population from 
Alto de Cuevas in Antioquia, Colombia, has by far the largest leaves of any 
population of this species known thus iar. Further study oi the population 
biologies of Cybunithus montciniis ( Lundell) G. Agostini from Panama, C 
oaigranatensis, and C. timanae will be necessary to fully resolve the precise 
relationships and microecological roles each plays in montane wet and plu- 
vial forests. 

4. Cybianthus spichigeri Pipoly, Candollea 46:43. 1991- (Fig. 2B, 1 1). 

Typf.: PERU. LoREit); Prov. Rec]uena, Trocha al Ajuajal, 2 km Irom Centro Eorestal 
Jenaro Jerrera, right bank of Rfo Ucayali, I 5 Eeb 1982 (stam. fl, tr), \i. Sptchiger & E. 
Encarniic'iihi 1224 (hoi.otypi;: US; is()rvpi;s: AMAZ, G, MO). 

Tree to 1 5 m tall. BraHchlets thin, straight, terete, 2—3 mm diam., densely 
lepidote. Leaves alternate; blades chartaceous, elliptic to narrowly oblanceolate, 
( 1 0— )1 5—20 cm long, (3— )5.2— 6.5(— 7.2) cm wide, apically caudate-acumi- 
nate, the actmien 1 .9—2.3 cm long, basally acutish to obtuse, not decurrent 
on the petiole, bullate, the midrib and secondary veuis strongly impressed 
above, prominently raised below, smooth and inconspicuously to promiently 
pellucid punctate above, moderately lepidote below, the margin essentially 
flat, but very slightly inroUed at the very margin; petioles canaliculate, (I—) 
1.2—2 cm long, densely lepidote. Staminate inflorescence: a pyramidal pinnate 
pannicle, 1-4.5 cm long, 1-3 cm wide, peduncle 0.3-1 cm long; branch 
bracts chartaceous, linear-subulate, 0.6—1 mm long, 0.1—0.2 mm wide, apically 
attenuate, densely lepidote; pedicels cylindrical, (0.8— )1— 1 .5 mm long. Staminate 
flowers 4— 5-merous, carnose; calyx suburceolate, 1.3—1.5 mm long, the tube 
0.3—0.5 mm long, the lobes deltate, ca. 1 mm long and wide, apically actite. 

PiPOLY, Cybianthus in Ecuador and Peru 


F](;. 11. Cybianthus spichigeri Pipoly. A. Habit, showin^q minute inflorescences and large, 
costate fruits. B. Staminate flower in bud, showing suburceolate calyx and tubiform co- 
rolla. C. Staminate flower, showing long, prominently lobate staminal tube, cuculiate co- 
rolla lobes, proximally recurved anthers. A-C, drawn from holotype, by Peggy Duke. Fig- 
ure reproduced from Pipoly, 1991- 

38 SiDA 1S(I) 

sparsely lepidote, prominently ru^M)se, with one prominent brown pancta- 
tion per lobe, the margins irregular, entire, sparsely lepidote; corolla tubiform, 
2.4—2.8 mm long, the tube ca. 0.5 mm long, the lobes oblong, 1.9-2.9 
mm long, 0.9-1.1 mm wide, apically acute, prominently cucuUate, with 
only a few, scattered scales without, glandular-granulose within, the mar- 
gin entire, glandular-granulose; stamens 2.2-2.4 mm long, the tube carnose, 
conspicuous, 1—1.4 mm long, lobate, the lobes 0.1-0.2 mm long alternat- 
ing with the filaments, the apex of the tube and lobes punctate, the apically 
free filaments, 0.9-1 3 mm long, the anthers deltate, 0.5-0.7 mm long and 
wide, apically apiculate, the apiculum slightly proximally recurved, basally 
cordate, the connective dorsally punctate with small brown dots forming a 
triangle along connective margin; pistillode conic, 0.5—0.7 mm long, 0.2— 
0.3 mm wide, hollow, glabrous, the style conspicuously brown punctate, 
the stigma punctiform. Pistillate and poly^ai//.()//s inflorescence: a raceme, occa- 
sionally a poorly formed panicle of 1—3 racemes branched from base, 0.4— 
1.5 cm long, densely lepidote, tardily glabrescent; peduncle 0.1-0.4 mm 
long; floral bracts chartaceous, ovate, 0.8-1 mm long, 0.3-0.5 mm wide, 
apically attenuate, densely lepidote; pedicels obconic, (0.8-)1.5-2.5(-3) mm 
long, to 1.2 mm diam. apically in fruit, densely lepidote. Pistillate flowers as 
in staminate but calyx 1.0-1.2 mm long, the tube 0.4-0.5 mm long, the 
lobes 0.6-0.7 mm long and wide, staminodes and ]Tistil unknown. Fr/nt 
depressed-globose, 0.7-0.8 cm long, 0.7-1.5 cm wide, prominently cos- 
tate longitudinally, the exocarp costate, inconspicuously pellucid punctate. 

Distribution. — Principally known from forests along the rfos Ucayali, Maranon 
and Napo Drainage Basin complex, Loreto, Peru, with one disjunct popu- 
lation in nearby Morona-Santiago, Ecuador, 150-1 80 (-1 ,500) m elevation. 

Ecology and conservation status. — Cybianthus spicbigeri is known from only 
a handful of specimens, but is locally quite common. Not enough is known 
of the population biology to categorize its conservation status, but its fre- 
quency in forest study plots of the Jenaro Herrera Reserve in Peru suggest 
it is reproducing and may not be in imminent danger. Cybianthus spicbigeri 
is a varillal or premontante sandstone species in Peru, and in Ecuador it is 
known only from premontane sandstones. Occurrence of this species in the 
Cerros del Sira, Peru, reinforces the concept that those mountains contain 
many unusual populations of otherwise lowland Amazonian plants. 

Etymology. — The epithet commemorates Rudolphe Spichiger, Director 
of the Conservatoire et Jardin Botaniques, Chambesy, Geneve, Switzerland. 
Dr. Spichiger has devoted much of his career to study of global change, 
conservation of biodiversity, and systematics of the genus Ilex. Under his 
leadership, the Jardin has maintained active research programs in Paraguay, 
Peru, Madagascar, and throughout Europe. 

PiPOLY, Cybianthus in Ecuador and Peru 39 

Representative specimens examined. ECUADOR. Morona-Santiago: Cordillera del Condor, 
Cuangos, 20 km E of Gualaquiza, near disputed Peru-Ecuador border, 03° 29' S, 78° 14' 
W, 1,500 m, 18 Jul 1993 (ster.), A. Gentry 80096 (BRIT, MO, QCNE), 1,470 m, 19 Jul 
1993 (infl. bud), A. Gentry 80179 (BRIT, MO, QCNE). PERU. Huanuco: Prov. Pachitea, 
region of Pucallpa, W part of "Sira Mountains," and adjacent lowland, ca. 24 km SE to 26 
km ESE of Puerto Inca, from beginning of rainforest to Campamento Pato Rojo, 09° 27' S, 
74° 46' W, 1,380 m, 31 Jan 1988 (fl bud), W. Moniwetz & B. Wallnbfer 14-31188 (BRIT, 
W, WU). Loreto: Prov. Maynas, Allpahuayo, IIAP Station, 04°10' S, 73°30' W, 150 m, 
13 Nov 1984 (fr), R. Vdsquez et al. 5911 (AMAZ, MO, NY), 6jun 1985 (fr), R. Vchquezet 
al. 6588 (AMAZ, MO, NY); Rio Nanay, Mishana, 30 km SW of Iquitos, 03° 55' S, 73° 
35' W, 150 m, 19 Aug 1978 (fr), R. Foster 4226 (MO, NY, USM), 16 May 1981 (fr), R. 
Va.u/uez &J. Crtollo 1801 (AMAZ, MO, NY), 20 Jan 1985 (fr), R. Vasquez & N.Jaramillo 
6/37 (AMAZ, MO, NY). Prov. Requena, Reserva Forestal JenaroHerrera, ()4°55' S, 73°45' 
W, along Rio Ucayali, 120 m, 1980 (stam. fl), R. Marimllod 9-R-137 (G, US), 1980 (bud), 
R. Manrnllod 4-R-90, 10 Jun. 1982 (fr), R. Sphhiner et al. 1973 (G, US), 24 Feb. 1987 
(ster.), A. Gentry et al. 5653 1 (AMAZ, MO); Aguajal, 3 km from Centro Forestal Jenaro 
Herrera, right margin Rfo Ucayali, 18 May 1982 (fr), F. Fncarnaaon 26105 (AMAZ, MO), 
22 May 1982 (fr), F Fncarnaaon 26200 (AMAZ, MO, NY, US); Arboretum, Centro For- 
estal Jenaro Herrera, 180 m, 13 Nov 1981 (fr, stam. fl), R. Spichiger & F. Fmarnacim 1027 
(AMAZ, G, MO, US). 

Cybianthus spichigeri is unique within subgenus Conomorpha because of its 
costate fruits. In addition, the subbuUate leaf blades, punctate staminal tube, 
obconic pedicels, and polygamous inflorescences are also exceedingly rare 
characters that allow for easy recognition. Since its description (Pipoly 1991), 
an entirely staminate specimen {R. Marmillod 9-R-137) has been located. 
While the staminate inflorescence structure is quite different from that of 
the polygamous one, flowers of both are identical. The occurrence of this 
taxon three times in one inventory conducted at the Jenaro Herrera Re- 
serve indicate that the relative frequency of reproductive individuals would 
permit a study of the breeding system and population biology for this most 
unusual taxon. 

5. Cybianthus lepidotus (Gleason) G. Agostini, Bol. Soc. Venez. Ci. Nat. 

22:388. 1976. Conomorpha lepidota Gleason, Bull. Torrey Bot. Club 58:146. 1931. 
Type: VENEZUELA. Tf.rritorio Fi^dhkal Ama/onas: Summit of Mt. Duida, 1,500 
m, Aug 1928-^Mar 1 929 (pist. fl), G. H. H. Tate 741 (hoi.otype: NY, F Neg. 040832; 
lsotype: US). 

Conomorpha curvii'enia Gleason, Bull. Torrey Bot. Club 58:444. 1931. Typp: VENEZU- 
ELA. Territorio Federal Amazonas: Mt. Duida, 1 ,260 m, Aug 1928-Mar 1929 (stam. 
fl), G. H. H. Fate 927 (holotype: NY; isotypi;: US). 

Conomorpha lepidota Gleason f. ac/itata Steyerm., Fieldiana, Bot. 28:465. 1953- Type: 
VENEZUELA. Territorio Federal Amazonas: Cerro Duida, 2 Dec 1 944 (fr),/. Steyermark 
58265 (holotype: F; isotype: NY). 

Shrub or small tree to 6 m tall. Branchlets straight, terete, 2-2.5 mm diam., 
densely lepidote. Leaves alternate; blades elliptic to narrowly elliptic, chartaceous 
to coriaceous, (3.1-)5-15 cm long, 1.6-6 cm wide, apically acuminate, the 

40 SiDA 18(1) 

acumen 0.3—2.0 cm long, basally acute to obtuse, midrib depressed above, 
prominently raised below, the secondary veins 24-28, inconspicuous above 
and below, pustulate and densely lepidote above at first, glabrescent, densely 
lepidote below, the scales not overlapping, inconspicuously pellucid punc- 
tate, the margin entire, subrevolute to revolute; petioles thin, marginate, 
1 .0-2.5 cm long, densely lepidote. S tarn mate inflorescence a spike, rarely two 
subsessile spikes, (2— )6-l 6 cm long, 8— 15-flowered, peduncle, pedicels, and 
axis densely lepidote, the scales not overlapping; peduncle (0.2-)0. 5-0.8 
mm long; floral bracts ovate to widely ovate, chartaceous, 0.5-0.6 mm long, 
0.3—0.4 mm wide, apically acute, densely lepidote adaxially, the margin 
entire, glabrous. Stcimincite flowers (4-)5-merous; calyx cupuliform, carnose, 
0.9-1.2 mm long, the tube 0.2-0.4 mm long, the lobes deltate to ovate- 
triangular, 0.6—0.8 mm long and wide, apically acute or acuminate, rarely 
obtuse, attenuate to a rounded tip, conspicuously brown punctate, the margin 
entire, lepidote; corolla cupuliform, carnose, 1.5-2.5 mm long, the tube 
0.5—1.0 mm long, the lobes ovate to broadly ovate, 1.2—1.7 mm long, 0.7- 
0.9 mm wide, apically attenuate to a round, cucullate tip, at times with a 
few, scattered lepidote scales without, glabrous without, glandular-granulose 
within over the entire surface, punctations brown, submarginal, the mar- 
gin entire, glabrous; stamens 1.5-1 .6 mm long, adnate 0.5-1.0 mm to corolla 
tube, the staminal tube 0.3—0.4 mm long, carnose, bearing lobes alternat- 
ing with the apically h-ee portions of the filaments 0.1-0.2 mm long, the 
filaments fiat, 0.3-0.4 mm long, erect, glabrous, the anthers dorsifixed less 
than 1/4 from base, ovate-triangular, 0.6—0.7 mm long, 0.4-0.5 mm wide, 
apically attenuate to a rounded tip, basally cordate, slightly dorsally reflexed, 
the connective epunctate; pistillode lageniform, 1.1-1.2 mm long, hollow, 
costate basally, sparingly translucent lepidote, pellucid-punctate. Pistillate 
inflorescence as in staminate but a spike, (2-)6-l6 cm long, 6-10-flowered; 
peduncle 0.6-0.8 mm long; fioral bracts 0.5-0.6 mm long, 0.3-0.4 mm 
wide. Pistillate floivers as in staminate, but staminodes 1.2—1.3 mm long, 
adnate 0.5-1 .0 mm to corolla tube, the staminodial tube 0.5-0.6 mm long, 
carnose, bearing lobes alternating with the apically free portions of the fila- 
ments 0. 1-0.2 mm long, those apical portions flat, 0.3-0.4 mm long, erect, 
glabrous, the antherodes dorsifixed less than 1/4 from base, deltate, 0.7- 
0.8 mm long and wide, apically attenuate to a rounded tip, basally cordate, 
slightly dorsally reflexed, the connective epunctate; pistil pyriform, 1.4- 
1.5 mm long, the ovary 1-1 .2 diam., the style not differentiated, the stigma 
punctiform, the placenta patelliform, bearing 2(-3) naked ovules. Fr/«V globose, 
purple at maturity, 0.5- 1.0 cm long, 0.6-1.0 cm diam., the endocarp smooth, 
the embryo curved, 3.5-4.0 mm long. 

Distribution. — Guayana Highland of Venezuela and Brazil, and sandstone 

PiPOi.Y, Cybianthus in Ecuador and Peru 41 

formations in Bolivia and Peru (reported for the first time here), 600—2,300 
m in Venezuela, 850-950 m in Bolivia, and 760-850 m in Peru. 

Ecology and conservation status. — Cybianthus lepidotm is restricted to large 
cloud forest formations in transition zones between sandstone and diabasic 
intrusions. It is often associated with species o^Erythroxylum, which are also 
edaphic endemics. It is a widespread, but locally infrequent species and therefore, 
should be considered threatened. 

Etymology. — The epithet refers to the densely lepidote vestiture of the 
vegetative and floral parts of the plant. 

Specimens examined. PERU. Amazonas; Prov. Bagua, Dtto. Imaza, Comunidad Aguaruna 
Putuim, anexo Yamayakat, SW of Putuim, 760-850 m, 26 Sep 1994 (stam. fl), C. Diaz et 
al. 7252 (BRIT, HUT, MO, USM). BOLIVIA. La Paz: Prov. Nor Yungas, valley of Rfo 
Coroico, Sacramento, 10 km NE of C^huspipaca on Coroico Rd., 27 Jan 1984 (seer.), A. 
Gentry & J. Solomon 44668 (MO), 4 km NE (above) Inabuara, 13.5 km above San Pedro, 
1,530-1,560 m, 22 Jan 1984 (stam. ti), A. Gentry &J. Solomon 44407 (MO); Prov. Larecaja, 
Maipiri, 6 Nov 1926-28 Feb 1 927 (stam. fl), 0. Buchtten 1 738 (HBG, NY, US), Copacabana, 
10 km S of Maipiri, 850-950 m, 8 Oct-1 5 Nov 1 939 (fr), B. Krnkoff 10987 (A, K, MICH, 
MO, NY, UC, US). 

Cybianthus lepidott/s, restricted to sizeable cloud forests in transition zones 
between sandstone and diabasic intrusions, is most easily recognized by its 
long, lax spikes. When sterile, it may be confused with Cybianthus roraimae 
(Steyerm.) G. Agost., but may be easily distinguished by the branchlets 2— 
2.5 (not 3.5—4.5) mm in diameter. It may also be confused with Cybianthus 
punctatus (Mez) G. Agost. and C. cardonae G. Agost. For a discussion of the 
differences between these taxa in sterile condition, see Pipoly (1992a). 

The area of Bolivia in which this species has been collected is of biogeo- 
graphic interest because it also supports several other Guayana Highland 
taxa in the Ericaceae and Clusiaceae. Its new discovery in Bagua Province 
of Amazonas, Peru, reinforces thematic map data from satellite imagery that 
indicated tepui-like vegetation could be expected in the area. It is interest- 
ing that C. Diaz et al. 7232 from Peru, and B. Krukoff 10987 from Bolivia, 
are qualitatively and quantitatively identical to specimens diCybianthus lepidotus 
from Cerro Duida, Amazonas, Venezuela. 

6. Cybianthus laetus (Mez ) G. Agostini (Fig. 2C), Acta Biol. Venez. 10:153. 
1980. Conomorpha laeta Mez m Engl., Pflanzenr. IV. 236(Heft 9):259. 1902. Type: 
PERU. Amazona.s: Taulia, without elevation or date (stam. fl.), A. Matthews ]y6l 
(hoi.otype: K; isotype: K). 

Shridb orstnalltree to 2 m tall. Branchlets straight, subterete, 2—3 mm diam., 
densely lepidote. Leaves alternate; blades coriaceous, obovate, 3-8(-12) cm 
long, 1.2-2(— 5.0) cm wide, apically acute or short-acuminate, basally cu- 
neate, decurrent on the petiole, midrib depressed above, prominently raised 
below, the secondary veins 14—18 pairs, planar above, conspicuous below, 

42 SiHA lcS(l) 

glabrous and smooth above at maturity, densely lepidote below, the mar- 
gin revolute; petioles marginate, 0.5—1 cm long, densely lepidote. Stami- 
nate inflorescence: a raceme or panicle with 2 racemose branches from base, 
2—3 cm long; rachis, and pedicels densely lepidote; peduncle ().1"{).3 cm 
long; pedicels cylindrical, 0.9—1 -5 mm long; floral bracts chartaceous, nar- 
rowly ovate, 1—2 mm long, densely lepidote adaxially. Staminate floiven 4- 
merons; calyx carnose, cotyliform, 1.2—1.8 mm long, sparsely to densely 
lepidote without, glabrous within, the tube 0.2-0.3 mm long, the lobes 
triangular to deltate, 0.8—1 mm long, 0.6—0.8 mm wide, apically attenu- 
ate to a rounded tip, inconspicuously pellucid punctate, the margin entire, 
lepidote; corolla carnose, campanulate, 3.2—3.4 mm long, the tube 1.8—2 
mm long, densely lepidote without, the scales overlapping, glabrous within, 
the lobes ovate or ovate-triangular, 0.8—0.9 mm long, 0.3—0.4 mm wide, 
symmetric, apically attenuate to a rounded tip, inconspicuosly pellucid punctate, 
densely lepidote without, glandular-granulose within, the margin entire, 
glandular-granulose; stamens 2.6—2.7 mm long, the tube carnose, conspicuous, 
equalling the corolla tube, 1.8-2 mm long, lobate, the lobes alternating 
with the filaments ca. 0.1 mm long, the apically free portions of the fila- 
ments flat, 0.1—0.2 mm long, the anthers dorsifixed, ovate, 0.6—0.9 mm 
long, 1.8—1.9 mm wide, somewhat curved distally, apically apiculate to 
subapicLilate, basally cordate, the connective inconspicuously brown punc- 
tate; pistillode conic, 1.5—2 mm long, translucent glandular-lepidote ba- 
sally, hollow. Pistillate inflorescence: as in staminate but 3—3.5 cm long; pe- 
dtmcle 0. 1-0.4 cm long; pedicels cylindrical, 0.9-1 .5 mm long; floral bracts 
chartaceous, linear-lanceolate, 1.3-1.5 mm long, densely lepidote adaxially. 
Pistillate flowers as in staminate but calyx 1 .2—1 .4 mm long, sparsely to densely 
lepidote without, glabrous within, the tube ca. 0.2 mm long, the lobes deltate, 
1 — 1.2 mm long and wide, apically attenuate to a rounded tip, inconspicu- 
ously pellucid punctate, the margin entire, lepidote; corolla, staminodes 
and pistil unknown. Fr//i/ globose, 5—8 mm long and indiam., exocarp carnose, 

Distribution. — Eastern slopes of the Andes, Colombia, Peru and Bolivia, 
1,980-2,850 m. 

Ecology and conservation status. — The species occurs in primary cloud for- 
est, a life zone being cleared rapidly for cultivation throughout the Andes, 
which may account for the paucity of collections. Within subgenus Conoviorpha, 
Cyhianthi/s laetiis is the species most in danger of extinction. 

Etymology. — The specific epithet is Latin for "cheerful or bright," and probably 
refers to the plant's aesthetically pleasmg appearance, having the same general 
form as many species of Waccinium, Myrsine dependens, other Ericaceae, and 
other diminutive Andean shrubs. The thick juicy exocarp is said to be very 
tasty although slightly acidic (T. Dudley, pers. comm.). 

PiPOLY, Cybianthus in Ecuador and Peru 43 

Representative specimens examined. COLOMBIA. Boyaca: Arcabuco, NE of town, 2,650 
m, 1 1 Nov 1965 (stam. fl), L. Uri/K s.n. (COL); Sierra Nevada del Cocuy, path from Laguna 
to Cobugon, near Alto del Oso, 2,900 m, 27 Aug 1958 (stam. fl), P. Gr/ihh et al. 744 (K). 
PERU. Amazonas: Prov. Luya, Dtto. Camporredondo, Anexo TuUanya, between Pajaco 
Tigreand Palma,06°04' 35"S,78° 21' 45" W, 2,500-2,600 m, 9 Dec I996(fr),J. Ciwpos 
et dl. 5UiI (BRIT, HUT, MO, USM); Along Rd. E of Chachapoyas between Pipos and 
Molinopampa, 06°15' S, 77°40' W, 1,980-2,340 m, 14 Feb 1985 (pist. fl, iv)J. Liiteyn & 
E. Cotton 11414 (NY, TEX, US, USM); E of Chachapoyas, 2,000 m, without date (stam. 
fl), A. Weberbaiier 43^4 (G). Cusco: Prov. La Convencion, Cordillera Vilcabamba, 12° 37' 
S, 73° 32' W, ceja and cumbre, 2,550 m, 3 Jul 1968 (pist. bud), T. Dudley 10690 (F, NA, 
USM), 5 Jul 1968 (pist. fl, fr), T. Dudley 10803 (F, NA, USM). Huanuco: Prov. Huanuco, 
Carpish Hills, trail to summit from W entrance, 2,700-2,850 m, 09° 42' S, 76° 05' W, 2 
Mar 1985 (stam. fl.), B. Stem & C. Todz/a 2292 (MO, USM). 

Cybianthus laet/ts is easily recognized by its small leaves, short petioles, 
subsessile anthers and thick, juicy exocarp. It is known outside of Peru from 
only two other collections, one from the department of Boyaca, Colombia 
and the other from Nor Yungas, Bolivia. A recent collection from Luya Province 
in western Peru (J. Campos et al. 3161) is referred here because of its thin, 
angulate branchlets, and the thick exocarp of its fruits, despite the larger, 
thinner leaves. It is expected in Ecuador, in either Napo or Santiago-Zamora 

7. Cybianthus peruvianus (A. DC. ) Miq. (Fig. 2D) in Mart., Fl. Bras. 

10:298. 1856. Conomorpba peruviana A. DC. , Ann. Sci. Nat., Hot. set 2, 16:92. 
\M\.Peckia peruviana (A. DC. ) Kuntze, Revis. Gen. PI. 402. 1891. Type: PERU. 
Amazonas: Moyobamba, 1838 (stam. fl), A. Mathews s.n. (holotype: G-DC; lsotypes: 
G, G-BOIS, GH, K). 

Conomorpha weherhaueri Mez, Repert. Spec. Nov. Regni Vcg. 3:101. 1906. Type: PERU. 
LoRirro: In mountains near Moyobamba, 1 ,300 m, 28 Aug 1904 (stam. fl), A. Weberbauer 
4668 (hoi.otypi:: B-destroyed; Lectotype, here designated: F). 

Shrub or small tree to 16 m tall. Branchlets straight, terete, 3-5 mm diam., 
densely lepidote. Leaves alternate, at times approaching pseudoverticiUate 
at some nodes; blades coriaceous, elliptic to obovate, 7-12.7 cm long, (2.6—) 
3.5—5 cm wide, apically acuminate, the acumen 1.2—1.6 cm long, basally 
acute to obtuse, decurrent on the petiole, midrib depressed above, promi- 
nently raised below, the secondary veins 10—25 pairs, prominently raised 
below, densely lepidote at first, then pusticulate above at maturity, sparsely 
lepidote below, the margin revolute; petioles canaliculate, 1—1.5 cm long, 
densely lepidote. Staminate inflorescence: a panicle with 1—4 racemes branch- 
ing from the base, (2— )3— 8 cm long; peduncle, rachis and pedicels densely 
lepidote; peduncle 0.1—0.2 cm long; floral bracts chartaceous, ovate, 0.7—1 
mm long, 0.5—0.6 mm wide, apically attenuate, densely lepdote abaxially; 
pedicels cylindrical, 0.5— 1(— 1.2) mm long. Staminate floivers 4— 5-mcrous; 
calyx green, cotyliform, 0.8—1 .2 mm long, lepidote without, glabrous within, 
the tube 0.3-0.4 mm long, the lobes ovate to deltate, 0.7—0.8 mm long, 

44 Sum 18(1) 

0.5—0.8 mm wide, apically artenuate ro a rounded tip, the margin lepi- 
dote, inconspicuously brown punctate; corolla translucent green, chartaceous, 
campanulate, 2—2.6 mm long, the tube 0.6— 1 .2 mm long, glabrous or sparsely 
lepidote externally, glabrous internally, the lobes ovate to narrowly ovate, 
1.2—1.8 mm long, O.")— 1.1 mm wide, asymmetric, apically rounded or at- 
tenuate to a roimded tip, sparsely lepidote externally, the scales not over- 
lapping, sparsely gland ular-granulose internally, inconspicuously brown punctate, 
the margin glandular-granulose; stamens 1 .4—2 mm long, the tube chartaceous, 
conspicuous, longer than the corolla tube, 0.8—1.2 mm long, elobate, the 
apically free portions of the filaments 0.2—0.4 mm long, the anthers linear- 
lanceolate, 0.7—0.9 mm long, 0.2—0.3 mm wide, somewhat recurved dis- 
tally, attenuate to a round or acute tip, the connective conspicuously brown 
punctate; pistiUode obclaviform, 1 — 1.5 mm long, translucent glandular- 
lepidote basally. Pistillate inflorescence: like the staminate but, 2—7 cm long; 
peduncle 0. 1-0.2 cm long; floral bracts 0.7-1 mm long, 0.4-0.5 mm wide; 
pedicels 0.2—0.5 mm long. Pistillate liowers as in staminate but calyx 0.9— 
1.1 mm long, the tube 0.3-0.7 mm long, the lobes deltate, 0.5-0.6 mm 
long, 0.4—0.5 mm wide, apically subacute to obtuse; corolla 1.3—1-7 mm 
long, the tube 0.2-0.3 mm long, the lobes ovate, 1-1.4 mm long, 0.7-0.9 
mm wide, apically obtuse; staminodes 1—1 .2 mm long, the tube longer than 
the corolla tube, 0.3—0.4 mm long, the apically free portions of the fila- 
ments 0.2—0.3 rnm long, the antherodes 0.6—0.7 mm long, 0,2—0.3 nim 
wide, pistil obnapiform, 1.3— 1.4mm long, 1-1.1 mm diam., the ovary 0.6— 
0.7 mm long, translucent glandular-lepidote basally, the style 0.5—0.6 mm 
long, conspicuously brown punctate, the stigma punctiform, the placenta 
globose, ovules 3, apically exposed. Fruit subglobose, 0.3—0.8 mm long, 
0.4—0.9 mm diam., the exocarp thin, black. 

Distribution. — Amazonian Ecuador through Peru to Bolivia, at 122—1,500 
m elevation. 

Ecology and conservation status. — Cyhianthus perui'ianus occurs on the east- 
ern slopes of rhe Andes in moist or wet lowland and premontane foresrs on 
white sands, especially in transition zones, where brownish sand-clay mix- 
tures occur. 

Etymology. — The epithet refers to the type locality, in (Moyobamba) Peru. 

Eocal navie. — Peru: "Tarrafactispi." (Quichua), "uchi yacushnum" (Aguaruna). 

Represencarivc sjiecimens examined. ECUADOR. Napo: CanttMi A^^tiarico, Reserva Fauni'stica 
Cuyabeno, Layima Zancutlo Cocha (Iripari), SE side ol Laguna, 00" 33' S, 75° 32' W, 230 
m, 28 Sep 1991 (fr), W. Palacios el at. 7761 (BRIT, MO, QCNH); Cant6n Orellana, Sector 
Huashito, 20 km N of Coca, PALMORIENTE property, 00" 20' S, 77° 05' W, 250 m, 3~ 
21 Nov 1989 (fr), E. Gi/uliiio 137 (BRIT, MO, QCNE); Sendero to Palma Roja, 28 Apr 
1986 (stam. ^)J.Jc/nimillo H522 (QCA). Zamora-Chinchipe: Canton Nangaritza Campamento 
Miazi, along Rfo Nangaritza, 900 m, 1 9 Eeb 1 994 (fr), //, van cler Werffet at. / J2<S'0 (BRIT, 
MO, QCNE); 11 ill above military post, 04" 18' S, 78° 40' W, 1,000 m, D. Nnll & W. 

PiPOLY, Cybianthus in Ecuador and Peru 45 

Palacios 9613 (BRIT, MO, QCNE), 04° 16' S, 78° 42' W, 970 m, 20 Oct 1991 (fr), W. 
Pa/acm eta/. 8486 (BRIT, COL, MO, QCNE). PERU. Amazonas: Along Ri'o Maranon, 
near confluence with Ri'o Santiago, 1924 (stam. fl.), G.>uann 3523 (B, G); Prov. Bagua, 
Dtto. Imaza, NE region of Rfo Maraiion Drainage Basin, Comunidad Kampaenza, along 
Qucbrada Shimutaz, Rio Maranon, 04° 55' S, 78° 19' W, .320 m, 9 Sep 1994 (pist. fl, Fr), 
N.Jaramilloetal. 436 (AMAZ, BRIT, HUT, MO, USM), 09 Oct 1995 (fr), N. Jaramillo & 
D. Chamik 813 (AMAZ, BRIT, MO, USM); Comunidad Aguaruna de Putuim (CAMPOU), 
anexo Yamayakat, Monte Alto de PutLiim, 450 m, 25 Aug 1994 (fr), C. D/az et al. 6993 
(BRIT, HUT, MO, USM), 660-760 m, 21 Sep 1994 (stam. fl), C, D/az et al. 7170 (BRIT, 
iMO, USM); Cerrosde Putuim, 05° O.V 20" S, 78° 20' 23" W, 350 m, 13Jun 1996 (fr),/?. 
Va'sqt/ez et al. 21131 . Huanuco: Prov. Pachitea, region of Pucallpa, W part of Sira Moun- 
tains and adjacent lowland, 20-24 km SE of Puerto Inca, Campamento Oro, 09° 29' S, 74° 
50' W, to Campamento Sira, 800 m, 17 Jul 1988 (stam. fl), W. Morawetz & B. Wallmfer 
22-19188 (BRIT, W, WU). Loreto: Prov. Loreto, Nauta, 04° 32' S, 73° 35' W, 160 m, 2 
Jun 1984 (fr), R. Vdsqiiez & N.Jaramillo 5060 (AMAZ, MO, USM); Ztingaru Cocha, 15 
km SE of Iquitos, 9 Sep 1964 (stam. fl), C. Dochon 2821 (AMAZ, MO, US, USM); Prov. 
Maynas, Moropon, lower Rio Nanay above Bellavista, 29 Aug 1968 (fr), S. McDaniel 1 0942 
(AMAZ, lEB, MO); Tamishiyacu, Quebrada Blanco Biol. Station, Camp II, Quebrada Blanco, 
Tahuayo River, 04° 23' S, 73° 17' W, 4 Apr 1 985 (fr),/. Castro 27 (AMAZ, MO, US, USM); 
Dtto. Iquitos, Hwy to Sto. Tomas, path in front ol "Chaparal" chicken farm, 140 m, 16 
Dec 1 983 (stam. fl), M. Rimachi 7232 (AMAZ, IBE, MO, US); Estacion Expetimental IIAP 
Allpahuayo, 21 km S of Iquitos,04° 10' S, 73° 30' W, 160 m, 18 Sep 1990 (ster.),/ P/poly 
etal. /2i72(AMAZ,MO,US, USM), 220 m, 19 Sep 1990 (stam. B) J. Pipoly et al. 12210 
(AMAZ, MO, NY, USM); Dtto. Sta. Maria de Nanay, 10 km W of Caserio Mishana, in 
Cocha Yarama Reserve, 03° 55' S, 73° 35' W, 130 m, 14 Mar 1991 (ster.),J. Pipoly etal. 
14994 (AMAZ, MO, US, USM), (ster.),/ Ptpalyetal. 14997 (AMAZ, MO, USM); Mishana, 
along Rfo Nanay, 03° SI'S, 73° 32' W, 150 m, 22 Apr 1986 (stam. bud), R. Vaujuez et al. 
7503 (AMAZ, MO, US, USM), 8 Sep 1990 (fr), R. Vdsquez etal. 14335 (AMAZ, BISH, F, 
MO, NY, TEX, US, USM); Mishana,() 3° 52' S, 73° 30' W, 140 m, 4 Jan 1983 (fr), A. 
Genttj etal. 39040 (AMAZ, MO, USM); Dtto. Iquitos, Puerto Almendras, Arboretum Ciencias 
de Ingenieria Forestal, UNAP, 03° 48' S, 73° 25' W, 122m, 4 Sep 1992 (ster.), C. Gra'/ulez 
et al. 471 1 (AMAZ, BRIT, MO, USM), Puetto Almendras, along Rio Nanay, 03° 45' S, 
73° 25' W, 122 m, 30 Oct 1984 (fr), R. Mchquez & N.Jaramillo 5867 (AMAZ, MO, NY, 
USM), 29 May 1986 (fr), R. Vazquez & N.Jaramillo 7570 (AMAZ, MO, US, USM), 30 
May 1986 (stam. bud), R. YdsqNez& N.Jaramillo 7587 (AMAZ, MO, US, USM); Quistococha, 
IMARPE tract, along Rd. 13 km from Iquitos, I4 Jul 1976 (fr), F. Encarnaaon 864 (AMAZ, 
US); Prov. Requena, Dtto. Saptiena, Jenaro Herrera, Rio Ucayali, 04° 55' S, 73° 40' W, 
160 m, 16 Aug 1994 (stam. fl), R. Ortiz et al. 98 (AMAZ, BRIT, MO, USM). Madre de 
Dios: Prov. Tambopata, Santuario Nacional Pampas del Heath, Quebtada Palma Real Grande, 
12° 57' 1 1" S, 68° 54' 48" W, 210 m, 21 Apr 1996 (fr), Al. Agu/lar & D. Castro 623 (BRIT, 
MO, USM), Puesto Enahuipa, 12° 39' 23" S, 68° 44' 13" W, 2 10 m, 25 Apr 1996 (fr), M. 
Agi/ilar & D. Castro 655 (BRIT, MO, USM). San Martin: Prov. San Martin, trail to televi- 
sion antenna, km 17.5 of Tarapoto-Yurimaguas Rd., 2.5 km N of Cataratas de Ahuashiyacu, 
06° 27' S, 76° 21' W, 850-1,200 m, 7 Sep 1986 (pist. fl, fr), S. Kuapp 8290 (MO, US, 

Cybianthus peruvianiis is most closely related to C. comperuviaJius Pipoly 
(herein described), but may be recognized by its longer, canaliculate peti- 
oles, revolute leaf margins, shorter pedicels, asymmetric, lepidote corolla 
lobes, and linear-lanceolate anthers. Field studies near Iquitos have shown 


SiDA 18(1) 

Fk,. 1 2. Cylnanthiii comperiivianus Pij^oly. A. Habit, showing inflorescence of racemes or malformed 
panicles. B. Staminate flower and axillanr bract, showint^ floral densely lepidote floral bract 
as long as pedicel, and lepidote calyx margins. C. Opened staminate flower, showing staminal 
rube longer than apical tree portions ot the lilamencs, lobes glandular-grannlose adaxially 
and crennlate margins. A--C, ch^awn Irom holotype, by Linny licagy. 

PiPOLY, Cybianthus in Ecuador and Peru 47 

it is a ridgetop species in the lowlands, with a density of approximately 2— 
4 individuals per hectare. 

8. Cybianthus comperuvianus Pipoly, sp. nov. (Fig. 2E, 12). Type: BRA- 
ZIL. Mato Grosso: Sta. Anna da Chapada, 1903 (stam. fl), G. Malme 3483 (holo- 
type: S; jsotypes: G, GH, R, S). 

Cybtanthus comperuvianus Agostini ex Pipoly in Killeen et al., Guia Arb. Boliv. 570. 1993. 
no?n. nud. Agostini (1972) provided the first description of this species in his disserta- 
tion, but never published it. I subsequently annotated herbarium specimens with the 
name, fully intending to publish it, but it was still not validly published when it 
appeared in Guia de Arholes de Bolivia, without Latin description or reference to type. 
Presumably, the name was obtained by them based on herbarium determinations, and 
thus a nomen nudum and invalid. Validation of the name is effected here,with the cita- 
tion of holotype above and the Latin diagnosis, and accompanying description, pro- 
vided below. 

Species haec cum C. peruviano saepenumero confusus est, sed laminis membranaceis (non 
coriaceis), secus margines planis (ncc revolutis), petiolis marginatis (non canaliculatis) 0.5- 
1 (nee 1-1.5) longis, pedicellis 1.5-2.5 (non 0.7-1.0) mm longis, coroUis staminaribus 
salverformibus (non campanulatis) statim diagnoscenda. 

Shrub or small tree to 6 m tall. Branchlets angulate, 1.5-2 mm diam., densely 
lepidote. Leaves alternate; membranaceous, elliptic, narrowly elliptic, to obovate, 
9-13(-17) cm long, 3-5(-6) cm wide, apically acuminate, basally acute, 
decurrent on the petiole, the midrib slightly depressed above, prominently 
raised below, pusticulate above, densely lepidote below, the secondary veins 
20-30 pairs, the margin flat; petioles marginate, 0.5-1 cm long, densely 
lepidote. Staminate inflorescence: a raceme, rarely a malformed panicle with 2 
branches from the base, 1-3 cm long; peduncle 0.1-0.5 cm long; floral bracts 
membranaceous, ovate, 1 .3-1 ■5(-2.0) mm long, ca. 0.6 mm wide, subglabrous, 
sparsely lepidote above and densely lepidote below, apically acute, slightly 
shorter than or as long as the pedicels, entire; pedicels cylindrical, 1.2-2.5 
mm long, sparsely lepidote. Staminate flowers whitish-green, 4(-5)-merous, 
membranaceous; calyx cupuliform, 1-1.2 mm long, the tube 0.3-0.6 mm 
long, the lobes triangular to deltate, 0.6-1 mm long, 0.5-0.7 mm wide, 
apically attenuate to a rounded tip, sparsely lepidote without, glabrous within, 
conspicuously brown punctate, the margin entire, sparsely ferrugineous- 
lepidote; corolla salverform, 2.2-2.6 mm long, the tube 1-1.2 mm long, 
the lobes ovate to ovate-triangular, 1.2-1 .6 mm long, 0.8-1 mm wide, apically 
rounded or attenuate to a rounded tip, glabrous without, glandular-granulose 
throughout within, the margin minutely crenulate, glandular-granulose, 
conspicuously brown punctate; stamens 1.8-2.2 mm long, adnate 1.2-1.5 
mm to corolla tube, staminal tube 1.6-1.9 mm long, elobate, longer than 
the apically free portions of the filaments 0.2-0.3 mm long, the anthers 
deltate to triangular, slightly distally curved, 0.7-0.8 mm long, 0.5-0.6 
mm wide, apically apiculate, basally broadly cordate, the connective incon- 

48 SiDA 18(1) 

spicuously brown punctate; pistillode lageniform, 1.0-1.2 mm long, 0.3— 
0.4 mm wide, the ovary 0.4—0.5 mm long, densely translucent glandular- 
lepidote near base, the style 0.5-0.6 mm long, the stigma punctiform. Pis- 
tillate inflorescence as in staminate but pedicel 1.5-2.5 mm long. Pistillate 
flowers as in staminate but calyx 1.0-1.2 mm long, the tube 0.3-0.6 mm 
long, the lobes deltate, 0.6-1 mm long and wide, corolla, staminodes and 
pistil unknown. Fruit globose, 5-7 mm long, 6-8 mm diam. 

Distribution. — Ecuador southward to Bolivia and adjacent Brazil, 400— 
1,200 m elevation. 

Ecology a)id conservation status. — Cybianthus coniperuvianus occurs in primary 
premontanc forests on sandstone and limestone, in relatively sparse popu- 
lations (teste coUectore). Therefore, it should be considered threatened. 

Etymology. — The specific epithet refers to the fact that Cybianthus comperuvianus 
has long been confused with C. peruvtanus. 

Local name. — Peru: "wewe" (Jivaro); "uchi apikna" (Huambisa); "CumaliUa" 

PARAT^-prs. ECUADOR. Morona-Santiago: Sitio La Planada, 01° 46' S, 77° 57' W, 
900 m, 2 1 Sep 1993 (stam. H), \V. Pciljcio.s 1 1407 (BRIT, MO, QCNE). PERU. Amazonas: 
Near Yucui Entsa, 6 hrs. from Camino tie Kusu, 300 m, 1 1 Mar 1973 (It), R. Kayap 538 
(MO, NY, USM), (fr), E. Ancuast) 93 (AM AZ, BRIT, F, LL-TEX, MO, NY, US, USM); Qucbrada 
Huampami, Tseasim, near Nayumpim, 244 m, 3 Apr 1973 (fr), E. Anaunlo 141 (AMAZ, 
F, LL-TFX, MO, US, USM); Quebracla Yurui Entsa, 305 m, 12 Apr 1973 (fr), E. Aiiauish 
220 (AxMAZ, F LL-TEX, MO, US); Prov. Bagua, Drto. Imaza, NE Region of Ri'o Marafion 
Drainage Basin, Comunidad de Yamayakac, Rio Maraiion, 04° 55' S, 78° 19' W, 320 m, 8 
Aug 1994(stam. H), N.Jaramillaetcil. 321 (BRIT, HUT, MO, USM). Ayacuclio: Tambillo, 
Toche Colorado, 27 Jul. 1878 (fr), C.Jelski 360 (PR, W). Huanuco: E of Tingo Man'a, 5 
Oct. 1972 (sram. (i), T. Croat 21194 (F, USM, MO); Agua Bianca, trail to Monzon, 9 Feb 
1966 (stam. 11.),/, Sitjmike 1049 (AMES, MO, NY, S, USM, VEN); vicinity Rondos, 24 
Mar. 1962 (sram. H.),_/. Sc/vnike 388! (F, US, USM); Prov. Pacliirea, region of Pucallpa, W 
part of Sira Mountains and adjacent lowland, 20-24 km SE of Puerto Inca, Campamento 
Oro, 09° 29' S, 74° 50' W, to C;ampamento Sini, 800 m, I "Jul 1988 (stam. fl), B. Walliwfer 
12-17788 (BRIT, MO, W, WU), 30 Aug 1988 (fr), W. Alonnmz & B. Wallnofer 13-30888 
(BRIT, W, WU), from Campamento Sira, 09° 28' S, 74° 47' W, SE to valley of Rio Negro, 
650 m, 1 I Aug 1988 (fr), W. Morawetz & B. Walinofer 1 13-1 1888 (BRIT, MO, W, WU). 
Juni'n: E of Quimiri Bridge, near La Merced, 800-1,300 m, I Mar 1929 (stam. H.), E. 
Killip & A. C. Simtlo 2401 1 (F, NY, US); La Merced, Hacienda Schunke, 27 Aug-1 Sep 
1923 (stam. fl.),/, Mac/mc/e5677 (F); Pichi's Trail, San Nicolas, 1 ,100 m, 4 Jul 1929 (stam. 
fl.), E. Killip & 'a. C. Smitl} 26(P3 (F, NY US), Sta, Rosa. 625-900 m, 6 Jul 1929 (stam. 
H.), 26768 (BM, F, NY, US, USM); Puerto Yessup, iOO m, 10 Jul 1929 (stam. fl.), /:. K//tip 
& A. C. Smitlj 26286 (NY, US); Puerto Bcrmudez, 375 m, 1 4 Jul 1929 (stam. H.), E Killip 
&A. C. Switlj 26464 (NY, US), 26548 (NY, US), 2656.1 (NY, US). Pasco: Prov. Oxapampa, 
Pichi's Valley, San Manas Ridge, 10-12 km SW of Puerto Bermudez, above Sta. Rosa de 
Chin's, trail to Loma Linda, 10" 20' S, 75° 00' W, 500 m, 29 Sep 1982 (fr), R. Eo.stcr et at. 
8962 (F, MO, USM). Puno: Below San Ciabon on Rfo San Ciabon, 500-1,000 m, 17-24 
Jul 1978 (stam. bud), Al. Dillomjtjl. 1219 (BRIT, F MO, LJSM). San Martin: Prov. Mariscal 
Caceres, Drto. Tocache Nuevo, Palo Blanco near Funtlo de Manuel Aranjo, 700-800 m, 1 

PiPOLY, Cybianthus in Ecuador and Peru 49 

Mar 1979 (fr) J. Schunk 10893 (AMAZ, BRIT, F, MO, NY, US); without specific locality, 
1778-1788 (stam. fl.), Ru/z L. &J. Patm 3/36 (F, MA), BOLIVIA. La Paz: Mapin Re- 
gion, 1926 (stam. fl.), 0. Buchnen 1733 (F, GH, HBG, NY, US); Tuiri, near Mapiri, 490- 
750 m, Sep 1939 (stam. fl.), B. Knikoff 10930 (A, G, GH, MO, MICH, NY, S, U, UC, US). 
Santa Cruz: Velasco; Parque Nacional Noel Kempff Mercado, Campamento las Gamas, 
14° 48' 41" S, 60° 23' 45" W, 850 m, 26 Mar 1993 (fr), L. Arroyo & K. Keill 164 (BRIT, 
MO, USZ); Campamento Huanchaca, 13° 54' S, 60" 48' W, 650 m, 17 May 1994 (stam. 
fl), L. Arroyo et ai. 61 4 (BRIT, MO, USZ). BRAZIL. Mato Grosso: Mpio. Cuiaba, Burity, 
NE of Cuiaba, 750 m, Jul 1927 (stam. fl.), B. Collenette 113 (NY); Chapada dos Guimraes, 
Cachoeira Veu de Noiva, do Rio Coxipozinho, 15° 30' S, 55° 45' W, 21 Oct. 1985 (fr),/ 
Prra>2! 1326 (INPA, MG, NY, SP), 720 m, 16 Oct 1973 (fr), G. Prance et a I. 19073 (AAU, 
F, INPA, K, MG, NY, K, S, SP, U, US); Sta. Anna da Chapada, 1902 (stam. fl), G. Malme 
2048 (S, UPS), 1903 (fr), G. Malme 3483 (G, GH, R, S), 1827 (stam. fl.), L. Riedel 939 
(LE, NY, US), 1902 (fr), A. Robert 322 (BM). Rondonia: 1 km NE of Ariquemes, Porto 
Vehlo-Cuiaba Hwy., 13 Aug 1968 (fr), £, Porero & L. Wrigley 7033 (MG, MO, NY). 

Cybianthus comperuvianus was first recognized as a novelty by Agostini (1972). 
Despite the fact that over 25 years has past since its first recognition, pistil- 
late flowers are still unknown, reinforcing the concept that within the ge- 
nus they are ephemeral (Pipoly 1983a, 1992). Cybianthus comperuvianus is 
most closely related to C. peruvianus but is easily distinguished by its long 
pedicels, salverform staminate corolla, and membranaceous leaves. 

9. Cybianthus guyanensis (A. DC.) Miq. in Mart, subsp. pseudoicacoreus 
(Miq. in Mart.) Pipoly, comb, et stat. nov. (Fig. 2F). Ardisia pseadoicacorea 
Miq. in Mart., Fl. Bras. 10:284. 1856. Conomorpha psendoicacorea (Miq. in Mart.) Mez 
in Engl., Pflanzenr. IV. 236(Heft 9):26l. 1902. Cybianthus pseudoicacoreus (Miq. in 
Mart.) G. Agostini, Acta Biol. Venez. 10:155. 1980. TYPE: BRAZIL. Amazonas: 
In forest near Rio Japura, Jan 1820 (pist. fl, fr), C. Martins s.n. (lectotype, here desig- 
nated: M; isolhctotype: M). 

Shrub or tree to 7 m tall. Branchlets terete, 2—3 mm diam., densley lepi- 
dote. Leaves alternate; blades membranaceous to chartaceous, narrowly obovate 
to elliptic, symmetric, (8-)9-12(-18.5) cm long, 3^.5(-5.7)cm wide, apically 
abruptly acuminate to caudate, the acumen 1-2 cm long, basally cuneate, 
decurrent on the petiole, midrib flat or slightly depressed on the upper surface, 
prominently raised below, the secondary veins 12-25 pairs, pustulate above, 
sparsely lepidote below, the margin flat; petioles canaliculate, 0.5— 0.8(— 1.5) 
cm long, densely lepidote. Staminate inflorescence: a raceme or panicle with 
1—2 branches from base, 1-3 cm long; peduncle, rachis and pedicels densely 
lepidote; peduncle 0.1-0.2 mm long; floral bracts chartaceous, narrowly 
ovate, longer than the pedicels, 1.3-1-5 mm long, 0.2-0.3 mm wide, apically 
attenuate, densely lepidote abaxially; pedicels cylindrical, 1—1.2 mm long. 
Staminate flowers 4-merous; calyx carnose, subcupuliform, 1—1.2 mm long, 
sparsely lepidote without, glabrous within, the tube 0.3—0.4 mm long, the 
lobes triangular, 0.8-l(-1.2) mm long, 0.5-0.7 mm wide, apically attenu- 
ate to a rounded tip, conspicuously brown ptmctate, the margin lepidote; 

50 SiDA 18(1) 

corolla carnose, camjoanulate, 2.8-3-2 mm long, the tube 1.2-1.8 mm long, 
glabrous, the lobes ovate, 1.3-1.8 mm long, 0.8-1 mm wide, erect to spreading, 
apically attenuate to a round tip, prominently or conspicously brown punctate, 
sparsely lepidote without, densely glandular-granulose on the upper half 
within, the margin entire, glandular-granulose; stamens 2.2-2.6 mm long, 
the staminal tube consj^icuous, carnose, 1.2—1.6 mm long, lobate, the lobes 
to 0.2 mm long, the apically free portions oi the filaments ().4-().6(-7) mm 
long, the anthers triangular, 0.7-0.9 mm long, 0.4-0.5 mm wide, slightly 
distally recurved, dehiscent by narrow introrse slits, apically attentuate to 
an apiculate tip, basally cordate, the connective inconspicuously brown punctate; 
pistillode elongate, conic, 1.2-1.8 mm long, densely translucent glandu- 
lar-lepidote basally. Pistillate inflorescence as in staminate but 1—3 cm long; 
lioral bracts 1—1 .3 mm long, ca. 0.2 mm wide; pedicels 0.6—0.9 mm long. 
Pistillate flowers as in staminate but calyx ca. 1 mm long, the tube 0. 1—0.2 
mm long, the lobes 0.8—1 mm long, 0.-1— 0.6 mm wide; corolla, staminodes 
and pistil unknown. Print subglobose, 4—6 mm long, 5—7 mm diam., exo- 
carp thin, pellucid punctate. 

Distribution. — Venezuela, Ecuador, Peru, Amazonian Brazil, and reported 
here for the first time from Bolivia, from 70—700 m. 

Ecology and conservation status. — Cybianthus guyanensis subsp. pseudoicacoreiis 
inhabits igapo forests of South-Central and Western Amazonia. It occurs 
in these forests on deep white sands just bek)w the floodline. It is periodi- 
cally inundated, but not for long periods. Quantitative fieldwork in Perti 
has shown it occurs in jiopulations of 8—1 individuals > 2.5 cm DBfi per 

Etymology. — The subspccific epithet refers to the growth habit of the plant, 
somewhat reminiscent oi Ardisia (subgenus I cacorea) guyanensis (Aublet) Mez. 

Representative specimens examined. ECUADOR. Napo: Canton Aguarico Reserva Etnica 
Huaorani, jVIaxus Oil Ilwy., km 60-61, S of Rfo Tivacuno, 00° 51' S, 76° 26' W, 250 m, 
21-25 Oct 1993 Or), /\l. A/ilcslia &J. Audi 923: Maxus Petroleum pipeline RcL, km 68, 
10 km SW of Ri'o Tivactino, 00" 49' S, 76" 2& W, 2fO m, 1 3 Dec 1993 (Ir), D. Matt a a/. 
10303 (BRIT, MO, QCNE). (BRIT, MO, QCNE); Estacion Experimental INIAP-Payamino, 
5 km NE of Coca, 00" 26' S, 77" 01' W, 250 m, 18-26 Feb 1986 (fr), W. Palmias d cil. 
I()4(> (MO, NY, QAME), (stam. fl), W. Palacios ct dl. lOSi) (MO, NY, QAME), 00° 25' S, 
77" 00' W, 250 m, 29 Nov 1 986 (fr), D. Nail 7494 (MO, QAME); Parqiie Nacional Yasiim', 
Lagunas de Garza Cocha, shore of Ri'o Garza, TO 1' S, 75 ' 17' W, 200 m, 22 Sep ! 997 
(fr), C. Ceroii &N. dllo "tOfij (MO, QCNE). trail hehind the house, 850 m, 27 Apr 1986 
{(v),J.Jdn/M/llo 8301 (QCA). PERU. Loreto: Prov. Maynas, Iquiros, G. Tessiiunin ,j65(; 
(NY), "l 00 m, 3-1 f Aug 1 929 (fr), II. Killip & A. C. Smith 2^005 (F, NY, US); Misluiyacu, 
near Iquitos, 100 m, 1930 (stam. il), G. Kl//;^ 1-412 (I'), I'eh 1932 (fr), G. Kl/zt^ 256^ (F, 
NY), 24 Sep 1 929 (stam. 11), I:. Killip & A. C. Smith 2<AS'" / ; Dtto. Alto Nanay, trail leading 
N from N end of Sta. Maria ck- Nanay, 5 Mar 1968 (stam. Ik), D. Simpson &J. Schittikt '^S4 
(F, US); Dtto. Indiana, Ex|ilorama Lodge, hallway between Indiana and mouth ol Rio Najio, 
03° 28' S, 72" 50' W, I sO m, 26 jun 1983 Or), /). Gt/itry et al. 42IH3 (AMAZ, MO), Far 

PiPOLY, Cybianthus in Ecuador and Peru 51 

end of Bushmascer Trail, l40 m, 5 Jan 1991 (ster.), A. Gentry et al. 12129 (AMAZ, MO, 
US, USM), Explorama Lodge, near Yanamono, 25 km NE of Iquitos, 03" 30' S, 72° 50' W, 
106 m, 24 Nov 1981 (fr), R. Vasquez & N. Jaramillo 2703 (AMAZ, MO, US, USM), Pe- 
rimeter path at southern boundary o^ reserve, 110 m, 27 Sep 1990 (ster.), J. Pipoly ct a!. 
12492 (AMAZ, MO, US, USM), 03" 28' S, 72° 52' W, 106 m, 15 Apr 1992 (fl bud), R. 
Vchq/wz & N.Jannmllo 18240 (AMAZ, BRIT, MO,USM); Dtto. Iquitos, Allpahuayo, Estacion 
Experimental del IIAP, 04° 10' S, 73° 30' W, 150-180 m, 3 Nov 1990 (fr), R. Vdsquez & 
N.Jarannllo 14545 (AMAZ, BRIT, MO, USM)8 Nov 1990 (ster.), R. Vasq//ez & N.Jarc/wilb 
15016 (AMAZ, MO, US, USM); Puerto Almendras, 2 Jul 1992 (fr), R. RimL, '&J. Ruiz 
597 (AMAZ, BRIT, MO); PuertoAlmendras, Arboretum Colegion Ingenerfa Forestal UNAP, 
03° 48' S, 73° 25' W, 122 m, 4 Sep 1992 (ster.), C. Grandez et cd. 4487 (AMAZ, BRIT, 
MO, USM); Dtto, Las Amazonas, Quebrada Sucusari, 03° 15' S, 72° 55' W, l40 m, 11 
Aug 1994 (stam. fl), R. Ortiz tt cd. 7-4 (AMAZ, BRIT, MO, USM); Dtto. Sta. Marfa de 
Nanay, Mishana, 03° 55' S, 73° 35' W, 90 m, 1 Ocr 1990 (ster.), J, Pipuly et al. 12706 
(AMAZ, MO, USM). Madre de Dies: Prov. Manu, Cerro de Pantiacolla, Rio Palotoa, 1 0- 
15 km NNW of Shintuya, 12" 35' S, 7 1 " 18' W, 650-700 m, 13 Dec 1985 (ster.), R. Poster 
et ill. 10993 (F, NY, USM); Prov. Tambopata, Santuario Nacional Pampas del Heath, Rio 
Heath, 12° 39' 23" S, 68° 44' 1 3" W, 2 1 m, 5 Jun 1 996 (stam. B), M. Aguilar & D. Castrn 
805 (BRIT, MO, USM). BOLIVIA. Santa Cruz: Dtto. Velasco, Parque Nacional Noel 
Kempff Mercado, Campamento La Torre, 13" 39' 20" S, 60° 49' 08" W, 200 m, 24 Nov 
1993 (fr), L Arroyo etcil. 510 (BRIT, MO, USZ). 

Cybianthus gNyanerisis s\.\hs\) . lisei/c/oiatcore/ts is one of three subspecies. The 
species is defined by the autapomorphic contorted anthers of the stamens 
and staminodes (Pipoly 1992a). The three subspecies may be separated in 
the following key. 

1. Leaf blades symmetric; calyx subcu|Tulilorm; corolla chartaceous or carnose, 
rhe lobes erect to spreading; apically free portions ot the filaments shorter 
than the anthers; anthers narrowly triangular or ovate-triangular, apically 
apiculate, dehiscent by narrow, introrse slits. 

2. Petioles canaliculate and winged, (1-)1 .5-1 .9(-2.3) cm long; staminate 
peduncle 0.2-0.5 cm long; floral bracts shorter than the pedicels, 0.7- 
0.8 mm long; pedicels 0.5 — 1 mm long; corolla chartaceous, 2.3—2.6 mm 
long, the lobes elliptic; staminal tube 0.9-1 ■ 1 mm long; apically Iree |X)rtions 

of the filaments 0.2-0.4 mm long subsp. guyancnsis 

2. Petioles canaliculate, 0.5-0.8(-1.5) cm long; staminate peduncle 0. 1-0.2 
cm long; floral bractslonger than the pedicels, 1 .3-1 .5 mm long; pedicels 
1-1.2 mm lonir; corolla carnose, 2.H-3.2 mm lont;, the lobes ovate; staminal 
tube 1.2—1.6 mm long; apically Iree |-)()rtions of the filaments 0.4—0.6 

(7) mm long subsji. pseudoicacoreus 

1. Leaf blades asymmetric; calyx cotylihirm; corolla membranaceous, the 
lobes reflexed-recurved; apically free portions ol the filaments longer than 
the anthers; anthers ovate, apically acute, dehiscent by wide, sublatrorse slits. 
subsp. multipunctatus 

Cybianthus subspecies multipunctatus (A. DC.) Pipoly is distributed in eastern 
Amazonia and the Guianas in premontane forests on lateritic and white sands 
of the Roraima Superimposed Sediments, while subsp. guyanensis is located 
principally in central Amazonia in igapo forests (Pipoly 1992a). Subspe- 


Si DA 18(1) 

Fic. 1 3. (.yhiciii!l.i/is tiDhDhiL l^ipoly. A. I labic, showint; paniculate iiiHoresccnces. B. Branchler 
apex. C. Adaxial leaf surlacc, showing midrib slightly raised but canaliculate. D. Abaxial 
leaf SLirhice, showing lepidote scales and prominently raised midrib. E. Section of raceme, 
showing coriaceous, delrate floral bracts, cotyliform calices, obcordate anthers with dis- 
tally recurved apiculae. F. Pistillate flower bud. G. Pistillate corolla with one lobe removed, 
showing lagenilorm pistil and siibsessile antherodes. A— F, drawn Irom holorype. F— (i, drawn 
Irom 'iinunsci l()4'^. Figure drawn h\ Finda Fllis. 

PfPOLY, Cybianthus in Ecuador and Peru 53 

c'les, pseudokacoreus is distributed south-central and western Amazonia, along 
banks of smaller streams on deep white sands. S\xhs]i&cies pseudokacoreiis and 
guyanensis are sympatric only in central Brazilian Amazonia, and differences 
in their ecology are not known in sufficient detail. In Ecuador and Peru, 
subspecies pseudokacoreus may be most easily confused with C comperuvianus 
Pipoly, but may be recognized by the thicker branchlets, longer floral bracts, 
and carnose perianth. 

10. Cybianthus timanae Pipoly, sp. nov. (Fig. 13). Type: PERU. Junin; Prov. Satipo, 
Gran Pajonal, Mapari, ca. 12 km SW of Chequitavo, 10° 45' S, 74° 23' W, 1,300 m, 
7 Apr 1984 (stam, fl), D, Smith 6782 (iiolotype: MO; isotypes: BRIT, US, USM). 

Propter ramulos graciles angulatos, laminas ellipticas vel oblanceolatas, ad apices acuminatas 
ad bases acutas, inflorescentiam paniculatam, calycem cotyliformem, necnon tubum staininarem 
staminodiaremque inconspicuo, C. occigranatens i a.Tcte affinis, sed ab ea nerviis secundariis 
16-40 (non 8-12)-]ugis, perianthiis chartaceis (non coriaceis), corolla cotylilormi (non 
campanulata), lobis corollinis laevibus (non verrucosis), ad apices acuminatis (nee attenuatis), 
conspicue (nee incounspicue) atro-punctaris, pistilo lageniformi (nee obnapiformi) pertacile 

Subshrub to 1 m tall. Branchlets angulate, 2-2.5 mm diam., densely lepi- 
dote. heaves alternate; blades chartaceous, elliptic to oblanceolate, (8—) 
9-5-1 5(-l 9) cm long, (2.7-)3-5.5(-6.5) cm wide, apically abruptly acuminate, 
caudate, the acumen 1—2.2 cm long, basally acute, decurrent on the peti- 
ole, smooth and nitid above, pallid and moderately lepidote below, midrib 
slightly raised and canaliculate above, not decurrent on the petiole, promi- 
nently raised below, secondary veins 16-40, brochidodromous, planar to 
somewhat impressed above, not bullate, the margin flat, entire; petioles 
canaliculate, (1.3-)1.5-2(-2.2) cm long, glabrous above, densely lepidote 
below. Staminate inflorescence: a panicle of 2—8 racemes branched from base, 
(3-)4-6(-7) cm long; peduncle 3-7 mm long; rachis densely lepidote; floral 
bracts coriaceous, deltate, 0.6-0.8 mm long and wide, apically acute, somewhat 
cucullate, densely lepidote above and below, the margin entire; pedicels 
cylindrical, 1-2.5 mm long, densely lepidote. Staminate flowers 4-merous, 
cream; calyx chartaceous, cotyliform, 0.7-1 mm long, the tube ca. 0.2-0.3 
mm long, the lobes ovate-triangular, 0.5-0.7 mm long, 0.3-0.4 mm wide, 
apically acute, moderately lepidote, the margin entire, somewhat involute; 
corolla chartaceous, cotyliform, 2—2.4 mm long, the tube ca. 0.2 mm long, 
the lobes oblanceolate, 1 .8—2.2 mm long, 0.8—1 mm wide, apically subacuminate, 
sparsely lepidote apically near margin, prominently black punctate and punctate- 
lineate without, densely glandular-granulose throughout within, the mar- 
gin glabrous, entire; stamens 1.4-1.6 mm long, the tube conspicuous, 
chartaceous, 0.1—0.2 mm long, elobate, the apically free portions of the filaments 
terete, 0.4-0.6 mm long, the anthers obcordate, 0.6-0.7 mm long, 0.3- 
0.4 mm wide, apically apiculate, anther and apiculum distally recurved, 

54 SioA LS(1) 

the connective prominently black punctate dorsally; pistillodc conic, 1.2— 
1.4 mm long, 0.3—0.4 mm wide, hollow, densely translucent glandular- 
lepidote. Pistillate inflorescence a^ in staminate, but (1— )1.5— 3 cm long; pe- 
duncle 2—4 mm long; floral bracts 0.4—0.6 mm long and wide; pedicels 
obconic, 0.6—0.9 mm long, sparsely iepiclote, conspicuously black ptmc- 
tate. PistilL/te flowers as in staminate but translucent green; calyx 0.7-0.9 
mm long, the tube ca. 0.2 mm long, the lobes 0. "3-0.7 mm long, 0.3—0.4 
mm wide, sparsely lepidote, corolla 1.2-1 .5 mm long, the tube ca. 0.2 mm 
long, the lobes oblong to oblanceolate, 0.9-1 -3 mm long, 0.4—0.6 mm wide, 
apically obtuse to subacuminate; staminodes 0.6-0.8 mm long, the tube 
conspicuous, chartaceotis, ca. 0. 1 mm long, the antherodes subsessile, 0.6— 
0.7 mm long, 0.3-0.4 mm wide; pistil lageniform, 1 .2-1.4 mm long, 0.3- 
0.4 mm wide, hollow, densely translucent glandular-lepidote, the ovules 
2-3, partially immersed on the placenta. Fr//il slightly depressed-globose, 
4.5—5.5 mm long, 5.5-7.5 mm diam., the exocar)-) thin, black at maturity. 

Distrih//ti())i. — Southeastern Ecuador to (4isco Peru, at 720—1,300 m 

Ecology and conservation status. — Cybianthiis t'nnanae usually occurs in wet 
premontane forest on sandstone soils. The restricted distribution oi Cyhianthus 
timanae indicates it should be considered a threatened species. 

Etymology. — The species is named for Martin Timana de la Flor, former 
Peruvian Field Associate ol the Missouri Botanical Garden, and currently a 
graduate student at the University of Texas at Austin. Martin is specializ- 
ing in the systematics of high altitude Caryophyllaceae. 

PARATYPF.S. ECUADOR. Zamora-Chinchipe: Nans^aruza Canton; lower slopes of 
Cordillera del Condor, above PachicLirza, Rfo Nangaritza Valley, 04° 07' S, 78" 3<S' W, 
1,000-1,200 m, 6 Dec 1990 (fr), D. Nell & W. Falacios 9536 (BRIT, MO, QCNB); Rio 
Nangarriza, Sliaime, confluence ol Rios Nangaritza and NLimpacakaime, 04° 20' S, 78° 
40' W, 1,000 m, 7 Dec 1990 (fr), /). Nail 9602 (BRIT, MO, QCNE); Pachicutza. Rd. to 
Hito, Cordillera del Condor, 04" 07' S, 78° 37' W, 1,000-1 ,100 m, 19 Oct 1991 (pist. fl 
bud), W. PciLuios i't cil. 8346 (BRIT, COL, MO, QCNE), 20 Oct 1 991 (stam. fl), W. Palaaos 
eta/. 8407 (BRIT, COL, MO, PORT, QCNE, USM); Parroc|uia Pachicutza, NE of military 
camp, 900 m, 6 I5ec 1990 (fr), _/. _/</;vw//^ & E. Cn/,/l,u/ 134/9 (COL, QCA). PERU. 
Cajamarca: CAitervo National Park, 1 2 km NE of San Andres de Cutervo, Transect 3, 06° 
10' S, 78" 40' W, 2,230 m, 10 Sep 1991 (ster.), A. G7;y/n c/ .//. 74630 (BRIT, MO, USM). 
Cusco: Pro\\ Quis|iicanchi. Camanci, Manin, along trail parallel to Ri'o Maniri to Quebrada 
Garrote, 13°I7'S, 70"48AV, 720 m, r Oct 1990 (pisr. fl), ,\I, T/m/i/a 1047 (BRIT, CUZ, 
MO, US, USM). 

Cybianthiis timanae is most closely relared to C occigranatensis (Cuatrec.) 
G. Agostini, but may be easily separated by its more numerous secondary 
veins, chartaceous perianth parrs, cotyliform corolla with smooth, conspicuously 
black punctate, actiminate lobes, and lageniform pistil. The involute calyx 
lobes and the obconic pedicels of the pistillate flowers are also extremely 
rare within the subgenus. 

PiPOLY, Cybianthus in Ecuador and Peru 


2 mm 



2 cm 

Fic;. 14. Cybianthus c/uitrecasasii Pipoly. A. Habit, showing irregularly shaped, basaliy branched 
panicles. B. Scaminate flower, habit, showing cotyliform calyx, infundibuliform corolla. C. 
Opened staminate flower, showing inconspicuous staminal tube and lageniform pistillode. 
D. Opened pistillate flower, showing obnapiform pistil and subsessile antherodes. A-C, 
drawn from holotype. D, drawn from A. Gentry et al. 33960. Figure drawn by Juan Pinzon. 

11. Cybianthus cuatrecasasii Pipoly (Fig. 14), Caldasia 18(3):285. 1996. 
Type: COLOiVIBlA. Huila: Municipio La Plata, Vereda Agua Bonita, Finca iVIerenberg, 
cerca del Cementerio (Jardi'n Botanico), 22 Abr 1982 (stam. fl)J. H. Torres 1034 

Dioecious tree to 6 m tall. Branchlets narrowly angulate, 3-4 mm diam., 

56 Sida18(1) 

densely appressed-lepidote, the scale margins frequently overlapping. Leaves 
alternate; blades chartaceous, elliptic or oblanceolare, 8—20 cm long, 3-5— 
6.5(— 6.9) cm wide, apically acuminate, the acumen (0.5-)l.()-1.5(-2.0) cm 
long, basally acute, slightly decurrent on the petiole, midrib impressed above, 
prominently raised below, the secondary veins 26-34 pairs, inconspicuous 
above, prominently raised below, sparsely lepidote above at first, then pus- 
tulate and glabrescent, densely appressed-lepidote below with margins fimbriate 
and overlapping, the blade margin flat, entire; petioles canaliculate, (1 .0)1 .3- 
1 .7(— 2.0) cm long, densely appressed-lepidote. Staniinate inflorescence: an irregular 
panicle with 2—4 basal branches, 4-8 cm long; peduncle, rachis and pedicels 
densely lepidote; peduncle 0.1—0.4 cm long; Horal bracts chartaceous, tri- 
angular, subequalling the pedicels, 1.2—1.5 mm long, ca. 0.6 mm wide, 
apically attentuate, densely lepidote below, the margin entire, glabrous; pedicels 
cylindric, 1 .0-1 .5 mm long. S tarn inate flowers 4-merous; calyx carnose, cotyliform, 
1.3—1.4 mm long, the tube 0.2—0.4 mm long, the lobes triangular or ovate- 
triangular, 1.0—1.2 mm long, 0.6-0.9 mm wide, apically attenuate, con- 
spicuously brown punctate, the margin entire, regular, with few scales; corolla 
chartaceous, infundibuliform, 3.0—3.5 mm long, the tube 1.2—1.3 mm long 
the lobes ovate, 2.0-2.2 mm long, 1.0—1.2 mm wide, apically rounded, 
carinate medially, densely lepidote without and along the margins, glan- 
duiar-granulose toward the apex and along the margins within, inconspicuously 
brown punctate, the margins regular, entire; stamens 2.5—3 mm long, the 
staminal tube membranaceous, inconspicuous, 1 .2-1 .3 mm long, hyaline, 
elobate, glabrous, the apically distinct hlaments adnate to the corolla lobe 
above the staminal tube 0.3-0.5 mm and apically free 0.1-0.3 mm, gla- 
brous, the anthers ovate, 0.8-1 .0 mm long, 0.5-0.6 mm wide, distally reflexed, 
apically apiculate, the apicuium slightly inHexed, basally cordate, the con- 
nective dark, eglandular; pistillode lageniform, 1 .6-1 .8 mm long, densely 
glandular-lepidote basally. Pistillate inllorescone: as in the staminate except 
a simple raceme, 4.4—10 cm long; peduncle 0.2-0.5 cm long; floral bracts 
lanceolate, 1 .2—1 .5 mm long, 0.3—0.4 mm wide; pedicels 0.7—1.3 mm long, 
incrassate in fruit. Pistillate flowers similar to the staminate except calyx 1 .4— 
1 .7 mm long, the tube 0.2—0.4 mm long, the lobes ovate, 1 .3—1.5 mm long, 
0.6-1.2 mm wide; corolla 3.5—4.0 mm long, the tube 0.9-1.1 mm long, 
the lobes elliptic, 2.7—3.1 mm long, 0.9-1 -3 nim wide, apically acute; staminodes 
similar to stamens but 3.0—3.2 mm long, the staminodial tube 1.0 mm 
long, the apically free portion of the filaments adnate to corolla lobes 1.3— 
1.5 mm and apically free ca. 0.1 mm, the antherodes subdeltate, 0.8-1.1 
mm long and wide, {always longer than wide), apically acute, basally sag- 
ittate; pistil obnapiform, 1.8-2.0 mm long, the ovary 0.9-1.0 mm long, 
1.2-1 .5 mm diam., densely translucent glandular-lepidote, the style 0.9- 
1 . 1 mm long, the stigma bilobate, not capitate, the jilacenta deeply cupuliform. 

PiPOLY, Cybianthus in Ecuador and Peru 57 

the ovules 2, deeply embedded in the placenta below apical pores. Fruit 
drupaceous, depressed-globose, 4-6 mm long, 6-8 mm diam., the endocarp 
smooth, the embryo erect, 3 mm long. 

Distribution. — Cybianthus cuatrecasasii occurs in the Western Cordillera 
of Colombia in the Department of Antioquia, in the Central Cordillera near 
the Macizo Colombiano, in the Departments of Huiia and Cauca, and in 
the Western Cordillera of adjacent Ecuador, with disjunct populations in 
the Province of Santiago-Zamora, and Zamora-Chinchipe, Ecuador, from 
(l,200-)l,960-2,850 m elevation. 

Ecology and conservation status. — Cybianthus cuatrecasasii occurs along the 
margins of primary cloud forests and along small watercourses. The popu- 
lation from Antioquia, Colombia is rare because it is from premontane plu- 
vial forest, and has leaves much shorter than normal. Because it is known 
only from primary forests, it should be considered threatened. 

Etymology. — ^This species is named to honor the late Jose Cuatrecasas, prodigious 
field botanist, monographer of many plant families, and ardent student of 
the Colombian flora. Don Jose freely offered his advice and assistance to all 
who asked, despite his limited time and numerous projects. 

Specimens examined. COLOMBIA. Antiocjuia: Mpio. San Luis, Piedra de Casrrillon, 
3-4 hours by foot S of town, 06° 01' N, 75° 01' W, 1,500-1,700 m, 8 May 1989 (A bud), 
D. Dalyetal. 3926 (HUA, MO, NY, US). Huila: Cordillera Central, E slope, Finca Merenberg, 
km 101 of La Plata-Popayan Rd., 13 km E ofSta. Letfcia, 02° 15' N, 76° 12' W, 2,300 m, 
24 Mar 1986 (fl bud), 6. Stein 3721 (BRIT, MO); Cordillera entre cuencasde los rios Guarapas 
y Guachicas, arriba de Palestma, al S\V de Pitalito, 2,000-2,300 m, 6 Feb 1943 (stam. fl), 
r.R. Pasherg 19969('NY, US); FincaMerenberg, E de Volcan Purace, ccrcade lazona limi'trofe 
con Cauca, 02° 26' N, 76° 12' W, 2,300 m, 1 Apr 1986 (bud), A. Gentry et al. 53881 
(COL, MO, US), 3 Apr 1986 (pist. fl, fr), A. Gentry et al. 53960 (COL, MO, US); 15 km 
NE de Algeciras, cerca del Campamento La Gironda, 2,400-2,850 m, 26 Mar 1944 (stam. 
fl)£. Uttle7481 (COL, NY, US);Mpio. Sanjosedelsnos, VeredaElHornito, 1,960-2,000 
m, 23 Jul 1980 (stam. fl), G. Lozano 3366 (COL); Mpio. La Argentina, arriba de Finca 
Palmira, 2,100 m, 26 Sep 1984 (fl bud), G. Lozauo et al. 4133 (COL). Cauca: Moscopan, 
margen del Rio San Jose, La Chorrera de Candelaria, 2,100-2,350 m, 1 Feb 1947 (fr),./. 
C//atrecasas 23583 (COL, F, NY, US); Cucnca del Rio La Plata, 43 km SE de Popayan, 2,1 60 
m, 26 Nov 1944 (stam. fl), F.R. Fosberg 22376 (NY, US). ECUADOR, Santiago-Zamora: 
along Quebrada Fionda, vicinity of Rancho Achupallas, 2,500-2,700 m, 1 Oct 1 943 (stam. 
fl),J. Steyermark 54552 (NY). Zamora-Chinchipe: Fiill ca. 2 km downstream from Campamento 
Shaime along Rio Nangaritza, 900 m, 15 Feb 1994 (fr), H. van ckr Werff 13074 (BRIT, 

Cybianthus cuatrescasasii is unique within subgenus Conomorpha because 
of its infundibuliform corolla and subapical portions of the filaments ad- 
nate to the corolla lobe. The overlapping covering scales of the abaxial leaf 
surface, branchlets and inflorescence rachis is found otherwise only in Cybianthus 
crotonoides (Mez) G. Agostini, a species endemic to the Guayana Highland. 
This species has been confused with Cybianthus peruvianus (Mez) G. Agostini, 

58 SiDA 18(1) 

but is easily separated from it by rhe infundibiiliform corolla, long pedicels 
and eglandiilar connectives. CybuDtthiis ciuitreccncisi'i is closely related to C 
Ictetiii (Mez) G. Agostini and C. oaigrciiiciteusis ((Aiatrec.) (3. Agostini, from 
which it is easily distinguished by its inftindibulitorm corolla, apical por- 
tions of the filaments adnate to the corolla and lack of lobes on the staminal 
and staminodial tribes. 

III. Cybianthus subgenus LaxifllorusG. Agostini, Acta Biol. Venez. 10:144. 

198{); Pipoly, Bnccoma 35:61-80. 1983. Cww^w^r/'^v secnon C<iiio)iii)ypbnhi W\l\., 
Stirp. Surinam Selc'c:. 111. Y'A'^Y^pyii pdvte. Coiioinorphc/ subgenus Eiaonouwrphu Mez 
in Engl., Pflanzenr. IV. 236(Heft 9):25-1 . 1 902. proper te nimore. Tyhi; Spiicii-s. Covoiiiorpha 
lax/Jlora (Mixrz.) A. DC. = Cybitnitlu/.s spiaiU/s {\\. B. K.)G. Agostini. (lecto'iype: by 
Agosrini, Acta Biol. Venez. 10:11'). 1980). 

Terrestrial dioecious shrubs or sniaU trees. Roots jiositively geotropic. Trunk 
distinguishable, leptocatiloiis, rhe growth dynamics following Rauh's Ar- 
chitectural Model (Halle et al. 1978). Brauehlets tomentose, with dendroid 
or stellate ferrugineous trichomes or both. Cataphylls and jiseudocataphylls 
absent. Leaves alternate, petiolate, tomentose, glabrescent. Inflorescence a simple 
raceme or rarely a spike, the peduncle 0.1-0.5 cm long, the staminate ra- 
chis tortuous or lax, the pistillate rigid and erect or rarely lax. Flowers A- or 
5-merous; calyx cotyliform, valvate, epunctate, inconspicuously pellucid or 
conspicuously brown or orange punctate, glabrous, tomentose, or glandu- 
lar-granulose, rarely bearing translucent glandular scales, the margins glandular- 
ciliate; corolla campanulare, rhe lobes valvate, glandular-granuiose only along 
margins without, but over the entire surface within, epunctate or incon- 
spicuously pellucid or conspicuously brown or orange punctate, and punc- 
tate-lineate, medially and basally glabrous, glandular-granuiose or with a 
few stellate trichomes; stamens and staminodes with filaments basally con- 
nate into a conspicuous or inconspicuous tube, the tube elobate or bearing 
small lobes alternate with the apically free filaments, the anthers elongate, 
triangular, distally curved, rarely erect, dorsifixed 1/4—1/2 from base, apically 
acute, basally cordate, dehiscent by wide longitutHnal slits; staminodes re- 
sembling sramens but reduced in size, the sterile anthers wirhout pollen or 
at times producing abortive pollen; pistil obnapiform, the ovary translu- 
cent glandular-lepidote, the style short, truncate with apunctiform stigma; 
pistillode lageniform, the ovary translucent glandular-lepidote, hollow the 
style elongate and curved apically, the nonfunctional stigma capitate. Fn/it 
drupaceous, one-seeded, the exocarp thin. 

Cybianthus subgenus Laxifloriis contains 6 species, 3 of which have been 
recorded from Peru. To date, none are known from Ecuador, but we may 
expect Cybianthus spicatiis (Kunth) G. Agostini in sandstone are^LS in the Cordillera 
del CcSndor area. 

PiPOLY, Cybianrhus in Ecuador and Peru 59 


1. Inflorescence rachis black |-iunctate; calyx lobes coriaceous, delcace, medi- 
ally carinate, basally rugose; fruit depressed-^ulobose, broader than long; large 
trees to 18 m tall, terra finne "varillal seco" [dry, on white sand] forests. 

12. C. nestor i i 

I. Inflorescence rachis epunctate; calyx lobes chartaceous, widely to narrowly 
ovate-triangular, medially thicked but flat; basally smooth; fruit globose, as 
broad as long; shrubs or small trees to 1()(-1 2) m tall; riparian gallery for- 
ests, either periodically flooded (varzea or iga|i6) or "varillal humedo" {wet, 
white or brown sand] forests. 

2. Leaf blades chartaceous, the tertiary veins prominently raised above; ca- 
lyx lobes longer than broad; staminal and staminodial tubes inconspicu- 
ous; receptacle or pedicel or both enlgarged in fruit; seasonally inundated 

forests (varzea or igapo) 1 3. C. spicatus 

2. Leaf blades coriaceous, the tertiary veins not visible from above; calyx 
lobes broader than long; staminal and staminodial tubes consjiicuous; 
neither receptacle nor pedicel enlarged in fruit; forests with 

poor drainage on sand, but non-intmdating, ("varillal humedo") 

M. C. fulvopulverulentus subsp. magnoliifolius 

12. Cybianthus nestorii Pipoly, sp. nov. (Fig. 15). Type: PERU. Lorpto: Prov. 
Maynas, Mishana, on Rfo Nanay, 03° 55' S, 73" 35' W, 150 m, 20 Jan 1985, 
R. Vcfsqmz & N. JaniwilU, 6122 (iiolotypi,: MO; istrtYPEs: AMAZ, F, NY, LJS, USM). 

Ob folia coriacea, rhachides inflorescentiares atro-punctato-lineatos, lobis calycinis deltatos 
inter se aequilatos coriaceosque, C. Jeltato valde arete afhnis sed ab ea habitu arboreo (non 
fruticoso) usque ad 18 (non 3) m, laminis anguste ellipticis vel anguste oblanceolatis vel 
anguste oblongis (non obovatis) 9.5-1 8(-28), (nee 3.2-4,4) cm longis, 3.2-4.6 (nee 1.7- 
2.8) cm latis, inflorescentiis 5-7 (non 1-4.2) cm longis, lobis calycinis 1.8-2 (non 1.3- 
1.7) mm longis, carinatisque (nee planisque) denique secus marginem minute erosis (nee 
undulatis), statim separabilis. 

Tree to 18 m tall. Branchlets terete, 5-7(-15) mm diam., appressed fer- 
rtigineous dendroid tomentose. Leaves alternate; blades thinly coriaceous, 
narrowly elliptic to narrowly oblanceolate, rarely narrowly oblong, (9-5-) 
11.5-14(-18) cm long, (3.2-)4-6 cm wide, apically obtuse, emarginate, 
rounded or acute, basally acute to attenuate, decurrent on the petiole, gla- 
brous and nitid above, glabrous and nitid but pallid below, midrib slightly 
raised above, prominently raised below, decurrent through length of peti- 
ole above and below, the secondary veins (l4-)20-47 pairs, prominulous 
(slightly raised) above and below, inconspicuously pellucid punctate and 
punctate lineate below, the margin entire, glabrous, revolute; petioles mar- 
ginate, (2.2— )2.5— 3.2(— 3.7) cm long, glabrous. Staminate inflorescence: un- 
known. Pistillate inflorescence: an erect raceme, 5-7 cm long, the rachis black 
punctate-lineate, ferrugineous glandular-granulose, glabrescent; floral bracts 
coriaceous, linear, minute, 1-1.1 mm long, 0.2-0.3 mm wide, apically acute, 
densely glandular-granulose below, glabrescent, the margin glandular-cili- 


SioA 18(i) 

Fk;. 1 5. Cyhuinthiii nestorii Pipoly. A. Habir, showini; depressed-^ lobosc fruits and leaf blades 
with prominuloLis secondary veins and revokice margins. B. Portion of infructescence, showing 
depressed-globose fruit and carinate, rugose calyx lobes with erose, sparsely glandular-ciliolate 
margins. (;. Abaxial leaf stirface, showing prominently raised midrib and promintilotis secondary 
veins. D. Branchlet apex, showing appressed dendroid tomentum. A-D, drawn from holo- 
type, by Linda Ellis. 

PiPOLY, Cybianthus in Ecuador and Peru 61 

ate; fruiting pedicels cylindrical, 2—4 mm long, densely glandular-granulose, 
minutely black punctate apically. Pistillate flowers unknown; fruiting calyx 
coriaceous, 1.8—2 mm long, the tube 0.3—0.5 mm long, the lobes deltate, 
1.3—1 .6 mm long, 1 .4—1.7 mm wide, apically acute, medially carintae, rugose 
basally, the margin minutely erose, sparsely glandular-ciliolate; corolla, staminodes 
and pistil unknown. Fr/zzV depressed-globose, 4-6 mm long, 6-8 mm diam., 
black at maturity, inconspicuously pellucid punctate. 

Distribution. — Known only from the Rio Nanay Drainage Basin, Maynas 
Province, Department of Loreto, Peru, at 150 m elevation. 

Ecology and conservation status. — Cybianthus nestorii occurs in "varillal seco" 
habitats, consisting of tall terra firme (non-inudating), relatively dry forest 
on deep white sands. It is a rare species and thus, should be considered threatened. 

Etymology. — It is a pleasure to dedicate this species to Nestor Jaramillo, 
of Iquitos, Peru, prodigious collector, and authority on plant collecting in 
tropical forests. Nestor, with his supervisor, Rodolfo Vasquez, form one of 
the most formidable botanical exploration teams in South American botany. 

Paratypes: PERU. Loreto: Prov. Maynas, Dtco. Iquitos, Puerto Almendras, UNAP, Tree 
No. 324, Tree inventory, 03° 48' S, 73° 25' W, 122 m, 17 Jan 1993 (ster.), C. Grdndez. N. 
Jaramil/o et al. 5321 (BRIT, MO, UNAP), Tree No. 373 (ster.), C. Grdndez, N.Jaramtllo et 
al. ')570 (BRIT, MO, UNAP, USM); Tree No. 651 (ster.), C. Grdndez. N. Jaramillo et al. 
5642 (BRIT, MO, UNAP). 

Cybianthus nestorii is unique within the subgenus because of its autapomorphic 
depressed-globose fruits and carinate calyx lobes. Other characters which 
readily seaparate it from its closest relative, Cybianthus deltatus Pipoly, of 
the Rio Guainia drainage basin of Venezuela, include its much larger ar- 
borescent habit, larger elliptic to narrowly oblanceolate leaves, longer 
inflorescences, and longer, minutely erose calyx lobes. 

13- Cybianthus spicatus (Kunth) G. Agostini (Fig. 3B), Acta Biol. Venez. 

10: 146. 1980. Mymm ipicata Kunth in H.B.K., Nov. Gen. Sp. 3:250. 1818. Omomorpha 
spicata (Kunth) Me2 in Engl., Pflanzenr. IV. 236(Heft 9):259. 1902. Tvpi;: VEN- 
EZUELA. Thiuutorio Fp.DBRAL Amazonas: In inundated area along the Ri'o Atabapo, 
without date (stam. fl), A. von Humboldt & A. Bonpland 1096 (iiolotypi;: P-BON; 
isotyph: B-destr.). 

Wallenui laxiflora Mart., Nov. Gen. Sp. PI. 3:89- 1829- Conomorpha laxiflora (Mart.) A. 

DC, Trans. Linn. Soc. London, Bot. 17:102. 1834, Typp: BRAZIL. Amazonas: Prov. 

Rio Negro, "In sylvis Japurensibus," Jan 1826 (stam. fl), C. Martins s.)i. (i.i;c:tot^'fi; 

by Pipoly 1983b: M; isolhctotype: CGE). 
Conomorpha laxiflora var. longifolia Miq. in Mart., Fl. Bras. 10:302. 1856. TvFi;: BRA- 
ZIL. Amazoxas: Prov. Rio Negro, vicinity Barra [Manaos], Dec-Mar 1850-5 1 (stam. 

fi), R. Spruce !()4() (LEf;i()TYPi"., here designated: M; isoleci-otypp.s, CGE, GH, GOET, 

K, LD, LE, OXF, U). 
Conomorpha laxiflora var. latijolia Miq. in Mart., Fl. Bras. 10:303. 1 856. Conomorpha tatijolia 

(Miq. in Mart.) Mez in Engl., Pflanzenr. IV. 236(HeIt 9):255. I 902. Typi;: VENEZUELA. 

62 SiDA 18(1) 

[CUYANA]. 1839 (scam. H), R. Sdmiiihiiriik 1002 (uicrorvPH by Pipoly 1983b: G; 

isoi.EC'iorYPr,: C^GE). 
Conomorpha catululleciiui Mcz in En^M., Pflanzenr. IV. 236(Heft 9): 23(3. 1 902. Typi': VEN- 
EZUELA. [GUYANA}. 1840 (scam, fl), R. Schowhuriik 88') (hopotypi:: B-dcstroyed; 

fragment at F; li-,c:ix)TYPi;, here clesignatetl: G; isoleci'otypi'.s: CGE, GH, K-2 .sheets, 

US, W). 
CdJKiiuorlibci grcitultjlorci Mez in Engl., Pflanzenr. IV. 236(HeI't 9):258. 1902. Typp.: 

HRAZIL.Amazonas: Rio Negro, above Moureira, Dec 1 85 1 (stam. fl.), R. Spr/ia- 1946 

(iioloiypp: B- destroyed; i.HC-roTYPP,, here designated: K; i.s()i.i;( totypps: CCtE, GH, 

Coimiumfbciglam-onihoisMfi m Engl., Pflanzenr. IV. 236(HeFt 9):260. 1 902. Type: BRAZIL. 

Ama/()\'as: Rio Negro, about 00° 30' S, (ri° 00' W, 21 Jun 18"?" 4 (stam. fl), J. Trail 

)()8 (hoi.otypp,: K). 
Coi/ijworplh/ >/k/(/c/m/si.\ A.C Smith, J. Arnold Arbor. 20:300. 1 93 I . Typi;: BRAZIL. Ama/.oxas: 

Mtinicipality oi: F^tmiayta, between Monte Cristo and Sta. Victoria on Rio Ipixtma, 

1 5-17 Nov 1934 (stam, fl), B. Krukoff724U (holotype: NY; isotypics: A, R M, MAD- 

Y, MICH, MO, S, US). 
Co>/oworp/ja gnni/is A.G. Smith, Bull. Torrey Bot. Glub 6'7:295. 1940. Tyi>i:: GUYANA. 

Artibaru River, Kako Territory, Upjier Mazaruni drainage, 600 m, 2 Feb 1939 (stam. 

fl), A. Pnikiis 181 (hou)type: NY; i,s()typi;s: BR, F, GH, M, MO, NY, S, US). 

Shi7/b or sijiall tree to 12m tall. Braiichlets thin to moderately thick, (.^.5-) 
4-8(-20) mm diam., appressed to floccose-dendroid and stellate ferrugin- 
eous tomentose, the stellate hairs often appearing lurfuraceous, glabrescent. 
Leaves alternate; blades tliinly coriaceous to chartaceous, to membranaceous, 
ovate, elliptic, obovate or rarely oblanceolate, (4.5— )6— 17.5(— 2 1 ) cm long, 
2.9— iS(— 1 1 .5) cm wide, apically actite, rounded, obtuse or rarely emargin- 
ate, basally acute to cuneate, glabrotis, pustulate and often nitid above, sparsely 
lerrugineous puberulent and minutely glandular-lepidote below, the mid- 
rib, secondary and tertiary veins prominently raised above and below, the 
midrib decurrent to the base of the petiole, pellucid or black punctations 
obscure below, the margin slightly revolute, entire; petioles distinctly marginate, 
(1— )1 .5— .3.2(— 4) cm long, stellate and dendroid ferrugineous tomentose, early 
glabrescent. Slciiiii)hite 'inflorescence, a simple raceme, lax, (3— )6-9(-l 2.5) cm 
long, the rachis epunctate, glandular-granuk)se with scattered stellate and 
dendroid ferrugineous trichomes, glabrate; floral bracts chartaceous, linear- 
lanceolate, ().7-L.7(-2.2) mm long, densely ferrugineous tomentose, early 
caducous; pedicels cylindrical, 0.9— 2.1 (-3) mm long, glabrous to densely 
glandular-granulose. Staimnate flowers (4— )5-merous, chartaceous, cream to 
yellow, with a sweet odor; calyx shallowly cotyliform, 1.5—2.5 mm long, 
the tube 0.3— 0.6 mm long, glandular-granulose at first, glabrescent, the 
lobes ovate or ovate-triangular, (1 .2-)l .4-2 mm long, 0.9—1.3 mm wide, 
acute to abruptly acuminate apically, somewhat cordate basally, prominently 
thickened medially, conspicuously punctate, the margins mostly entire, but 
occasionally erose apically, glandular-ciliate; corolla campanulate, 3.6—4.8 
mm long, the rube 1-1.2 mm long, the lobes ovate, 2. 1-3.4 mm long, 1.2— 

PiPOLY, Cybianthus in Ecuador and Peru 63 

2.6 mm wide, slightly reflexed at anthcsis, obtuse to acute apically, epunctate 
or conspicuously brown punctate; stamens 1.6—2.2 mm long, the tube 
membranaceous, inconspicuous, 1—1.5 mm long, the apically free portions 
of the filaments 0.3-0.55 mm long, the anthers elongate-triangular, 0.7- 
1 .2 mm long, 0.2—0.3 mm wide, dorsifixed ca. 1/3 from base, apically acute, 
basally cordate, distally recurved, the connective inconspicuously brown punctate; 
pistillode lageniform, 1.3-1.8 mm long, the ovary 0.6—0.9 mrn long, 0.5— 
0.75 mm diam., densely translucent glandular-lepidote, the style elongate, 
0.8-1.3 mm long, slightly curved apically, the stigma capitate, 0.1-0.2 mm 
long. Pistillate inflorescence as in staminate, erect, (3.3-)4.5-13(-17.5) cm 
long; floral bracts 0.5-2.4 mm long; pedicels 1.3-1.8 mm long, accrescent 
in fruit to 3-1 mm long, the receptacle or pedicel or both incrassate in fruit. 
Pistillate flowers as in staminate but chartaceous, yellow; calyx cotyliform, 
1.3—1-9 mm long, the tube 0.6—0.9 mm long, the lobes widely to narrowly 
ovate, 1.1-1.7 mm long, 0.9-1.5 mm wide, erect, sparsely brov/n punc- 
tate, prominently thickened below, the margins entire or occasionally erose 
apically; corolla 2.7—3.5 mm long, the tube 0.7—0.9 mm long, the lobes 
1.6—2.7 mm long; staminodes as in stamens but 1 .6—2.1 mm long, the tube 
chartaceous, conspicuous, 0.7-1.1 mm long, the apically free portions of 
the filaments 0.3-0.5 mm long, the antherodes 0.6-0.9 mm long, 0.2—0.3 
mm wide, at times producing abortive pollen; pistil obnapiform, 1.6—2.2 
mm long, the ovary 0.8-1.2 mm long, 0.9-1.2 mm diam., densely trans- 
lucent glandular-lepidote, the placenta cupuliform, ovules 4, partially im- 
bedded, the stigma capitate, 0.1-0.2 mm long. Fruit globose, 2.5-6 mm 
long and in diam., prominently black punctate-lineate, with a tew persis- 
tent lepidote glandular scales below the usually persistent style base, green 
then purple, then black. 

Distribution. — Venezuela, Colombia, Peru, Brazil and Guyana, in inun- 
dated forests (varzea and igapo), from 100—400 m. 

Ecology and conservation status. — Cybianthus spicatus is restricted to primary 
riparian habitats on white sandy soils. It is common in igapo and varzea, 
but the latter only when sufficient c^uantities of sand exist, a mixed varzea 
type. While it is a widespread species, it is sensitive to soil compaction, 
and should be considered threatened. 

Etymology. — The epithet refers to the inflorescence shape, which is a raceme 
bearing flowers on short pedicels, thus appearing spicate. 

Represenrarive specimens examined. PERU. Huanuco: Cerros del Sira, SW slope of 
the Rfo Lulla Pichis, 1 190 m, I 2 Jul. 1 969 (fr),_/, Wolffe 12295 (F, NA); 100 m, 22 Jul 
1969 (fr), 7.' D//cllty H 1 24 (F, NA); Prov. Pachicea, region of Pucallpa, W part of Sira Mountains 
and adjacent lowland, ca 24 km SE ro 26 km ESE of Puerto Inca, next to Campamento 
Pato Rojo, 09° 27' S, 74" 46' W, 1 ,000 m, 27 Jan 1 988 (pist. fl), W. Momwetz & B. Wallnofcr 
72-27788 (BRIT, MO, W,W[]), 1,320 m, 1 May 1988(fr),/3. WW/«r//tT 7 7 /-/.5<S\S' (BRIT, 
W, WU), 1 ,230 m, 1 3 Jun 1 988 (fr), B. Wcillndfer I I2-I368ti (BRIT, W, WU). San Martin: 

64 Si DA 18(1) 

Lamas, on old trail from San Antonio ck- Cumbasa, S oi Shapajilla, upper slopes of Cerro 
Isco, 06" 22' S, 76° 23' W, 6()()-S()0 m, 5 Oct 19H6 (btul), S. Knapp et al. 85 14. (fr), S. 
Kncipp €t al. 8") 11 (MO, US, USM). 

Cybtanthus spicatiis is a polymorphic ochlospecies, sensu White (1962), 
Prance (1972) and Pipoly (1983a), wirh many semi-isolated populations 
throughout the Amazon and eastern Guayana Floristic Province (Maguire 
1979). These localized populations have produced several seemingly dis- 
tinct ecotypes, resulting in overdescription. Collections oiCybianthus spkatiis 
from Peru match the type oi Conomorpha gracilis. A. C. Smith, described from 
Guyana, in every detail. Fieldwork conducted in Guyana, Venezuela, Bra- 
zil and Peru since my earlier treatment (Pipoly 1983a), where I discuss the 
four ecotypes present within the species, and the synonymy rationale, has 
confirmed the concept that Cyhianthus spicatns is a polymorphic ochlospecies. 

Steyermark (1981 ) discussed the fact that Richard Schomburgk's collec- 
tions attributed to Guyana are now known to be from Venezuela. 

14. Cyhianthus fulvopulverulentus (Mez)G. Agostini subsp. magnoliifolius 

(Mez) Pipoly (Fig. 3 A), Brittonia 35:72. 1983. Commwrpha magmliifolia 
Mez in Engl., Pflanzenr. IV. 236(Heft 9):258. 1902. Cybunithm »ui\^u,iliifdiin (Mez) 
G. Agostini, Acta Biol. Venez. 10:146. I 980. Type: SURINAME. without locality, 
(scam. II), H. Kt',v.e/ 244 (i.ectotyph by Pipoly 1983a: GOET). 

CoiniiiKirphi! rigicLi Mez, Repert. Spec. Nov. Regni Veg. 16:420. 1920. syn. nov. Type: 
BRAZIL. Amazona.s: "Hylaea," without date (Fr), E. Ilk 8722 (iiolotype: B-desrr., F 
Neg. 4831). Despite searches of herbaria housing significant Ule collections, no du- 
plicates of the type have been located. Therefore, I select a neotype herewith: Type: 
C;LJYANA [BRITISH GUIANA]. Orealla Savanna, Corenryne River, Sep 1879 (fr), 
/;. Imth/irn 6/9 (neotype here designated: K; KSONEOIyte: BRG). The neotype was annotated 
by Mez in 1901, bur was not mentioned in the protologue. 

Shr/ib or tree to 10 m tall. Branchlets mostly thick, (3-)5.2-12 mm diam., 
glabrate or densely stellate and dendroid ferrugineous tomentose and glan- 
dular-granulose, at times glabrescent. Leaves alternate, at times clustered 
and approaching pseudoverticillate; blades coriaceous, elliptic, oblanceolate, 
ovate or obovate, 7.2-21 .7 cm long, (2.2-)3.6-9.5(-17.1) cm wide, apically 
obtuse, acute, rounded or emarginate, basally obtuse to acute (rarely acuminate), 
sparsely pitted with superimposed glandular lepidote scales and glabrous 
or glandular-granulose, at times sparsely ferrugineous tomentose above and 
below, glabrescent, eptmctate, the margin entire and revolute; petioles marginate, 
(1 .3—)! .6— 4.2(— 4.6) cm long, ferrugineous tomentose, and glandular-granulose, 
glabrescent. Staininate inflorescence: a simple, tortuous raceme, rarely lax, (3.6-) 
4.5-1 5 cm long, the rachis maroon, opaque, glandular-granulose or with a 
few scattered dendroid ferrugineous trichomes, glabrescent; floral bracts 
chartaceous, linear-lanceolate, 0.8—3.6 mm long, ca. 0. 1—0.2 mm wide, densely 
ferrugineous tomentose, caducous; pedicels cylindrical, ({).2-)2.9— 3.6 mm 

PiPOLY, Cybianthus in Ecuador and Peru 65 

long, densely glandular-granulose, glabrescent. Sta77iinate flowers 4- or 5-merous, 
chartaceous, white, cream or yellow; calyx deeply cotyliform, 0.9-2.6 mm 
long, densely glandular-granulose then glabrescent, the tube 0.3-0.7 mm 
long, unequally divided, the lobes chartaceous, ovate to ovate-triangular, 
0.6-1.9 mm long, 1.1-2.4 mm wide, obtuse to acute or acuminate apically, 
prominently orange punctate, densely ferrugineous tomentose and ferrugi- 
neous glandular-granulose or partially glabrescent or ferrugineous glandu- 
lar-granulose and then sometimes totally glabrescent, the margins extremely 
undulate, entire, densely glandular-ciliate; corolla campanulate, (2.8-)3.3- 
4.7(_5.5) mm long, the tube (0.6-)0.9-1.4(-1.7) mm long, the lobes ovate, 
(2-)2.2-3.1(-4) mm long, (1.2-)1.5-2.4(-2.8) mm wide, the lobes highly 
reflexed at anthesis, obtuse to acute apically, conspicously orange punctate 
and punctate-lineate, at times scattered ferrugineous trichomes before an- 
thesis without, the margin entire; stamens (2.4-)2, 8-3-4 (-3.8) mm long, 
the staminal tube 0.8-1.1 mm long, the apically free portions of the fila- 
ments 0.6-0.9 mm long, the anthers elongate-triangular, (0.8-)l-1.4 mm 
long, apically acute, basally cordate, dorsifixed ca. 1/4 from base, distally 
recurved or rarely and aberrantly erect; pistillode (1.6-)1.9-2.5 mm long, 
the ovary 0.7-1.1 mm long, 0.7-1.2(-1.4) mm diam., densely translucent 
glandular-lepidote, the style elongate, curved, 1-1.6 mm long, the stigma 
0. 1-0.2 mm long. Pistillate inflorescence as in staminate but erect (3-)4-l 1 .4 
cm long, the rachis green then red; floral bracts 0.7-2.9 mm long, ca. 0.1- 
0.2 mm wide; pedicels cylindrical, (0.2-)0.6-1.5(-2) mm long. Pistillate 
flowers as in staminate but dull yellow to brown; calyx 1.2-2.6 mm long, 
the tube 0.4-0.7(-l) mm long, the lobes 0.7-2 mm long, 1 .3-2.4 mm wide; 
corolla (2.7-)3.2-3.5(-4) mm long, the tube 0.6-1.2 mm long, the lobes 
2-2.3(-2.6)mm long, 1.1-1.5 mm wide; staminodes as in stamens but 2.4- 
2.9 mm long, the staminodial tube 0.7-0.9 mm long, the apically free portions 
of the filaments 0.7 mm long, the antherodes triangular, 0.6-0.9 mm long; 
pistil obnapiform, 1. 9-2. 5 (-3) mm long, the ovary (0.7-)0. 9- 1.6 mm long, 
(0.7-)1.3-1.7(-2.2) mm diam., densely translucent glandular-lepidote, the 
style thick, (0.4-)0.6-0.9(-1.3) mm long, the stigma pseudocapitate, very 
minutely 4-lobed, up to 0.3 mm long. Fruit globose, 3-9 mm long and in 
diam., green, then red, then black, inconspicuously pellucid punctate. 

Distribution. — Venezuela, Guyana, Surinam, French Guiana, Brazil (Amazonas, 
Roraima, Para, Mato Grosso) and reported here for the first time from Peru 
and Bolivia 50-500(-850) m. 

Ecology and conservation status. — Cybianthus fulvopulverulentus subsp . magnoliifolius 
is extremely common in wet tepui savannas, but is rare in Peru in varillal 
humedo (wet sandy, non-inundating) habitats. It is anticipated, but has not 
yet been collected, in Ecuador. 

Etymology. — The epithet, 'fulvopulverulentus' refers to the rusty tomen- 

66 Si DA 18(1) 

turn of the species, forming a powdery like vestiture on ciie adaxial leaf surface, 
caducous as the leaf matures. The subspecific epithet refers to the shape and 
shiny adaxial leaf surface, giving the general impression of a Magnolia leaf. 

Specimens examined. PERU. Loreto: Prov. Maynas, Iquicos, Nma rumi-Ri'o Nanay, 03° 
48' S, 73" 25' W, 122 m, 5 Mar 1987 (bud), R. Vau/uez et al. 8905 (AMAZ, MO, US, 
USM). BOLIVIA. Santa Cruz: Prov. Velasco, Parque Nacional Noel Kempff Mercado, 
Campamenco Huanchaca I, 3.35 km from Rio Pauserna, 13° 56' 01" S, 60° 49' 30" W, 
600 m, 22 May 1994 (fr), L. Arroyo et al. 738 (BRIT, MO, USZ), 26 Jul 1995 (pist. H), T. 
Killeen et al. 7328 (BRIT, MO, USZ), (stam. ft), T. Ktlleen et al. 7330 (BRIT, MO, USZ), 
850 m, 3 Nov 1995 (Ir), A. Rodriguez &J. Suruhi 66'-) (BRIT, MO, USZ). 

Specimens cited here are the first known from Peru and Bolivia. Those 
collections represent remarkable disjunctions, the closest population known 
heretofore occurring approximately one degree north of the Rio Negro in 
southern Venezuela near the Brazilian frontier, or halfway across Brazil to 
the east, on Serra do Cachimbo, state of Mato Grosso. To date, no speci- 
mens are known from Ecuador. 

In my previous treatment (Pipoly 1983a), I included Conomorpha rigida 
in the synonymy oi Cybianthus fulvopulverulentus (Mez) G. Agostini subsp. 
fulvopulverulentm. The type at Berlin had been destroyed and I synonymized 
it based on the description and photo. However, the discovery of two du- 
plicates of the Imthurn gathering annotated by Mez permitted neorypification 
of the name with the Kew specimen and its inclusion here. Populations 
represented by this collection differ from the type of subspecies magnoliifolius 
only in their diminutive stature and vegetative parts. 

IV. Cybianthus subgenus Comomyrsine (Hook, f ) G. Agostini, Acta Biol. 

Venez. 1 0:162 1 980. Cuimmyrsme Hook. f. m Bench. & Hook., Gen. PL 2:643. 1 876; 
Wetgeltia A. DC. subgcniis Comomyrsine (Hook, f.) Mez in Engl., Pflanzenr. IV. 236(Hefc 
9):290. 1 902. Pax /« Engl. &Prantl, Nat. Pflanzenfam. IV, 1:92, 1897;J.F. Macbr., Field 
Mus. Nat. Hist., Bot. Ser. 1 3 (5, 1): 175. 1 959- Type Spi;cies: Cybianthus sprucei {Hook. 
f.) G. Agostini (m:ct()type: by D'Arcy, Ann. Missouri Bot. Gard. 60:445. 1973.) 

Terrestrial, erect, monoaxial suhshriibs or treelets to 2(-5) m tall. Roots positively 
geotropic. Bark mostly brown, smooth, longitudinally fissured or transversely 
checked. Trunk distinguishable, leptocaulous, monoaxial, following mor- 
phogenetic dynamics of Corner's Architectural Model (Halle et al. 1978), 
growth rythmic. Stems terete distally, sparingly to densely glandular-papil- 
late, at times with rufous hydropotes or orange glandular lepidote scales, 
without lenticels. Cataphylls alternate or pseudoverticillate, alternating with 
pseudoverticels of leaves, or apparently axillary to them, linear-subulate to 
acicular, rigid to membranous, keeled or flat, prominently punctate or punctate- 
lineate, glabrous, glandular-papillate, bearing hydropotes, or orange lepi- 
dote scales. Leaves pseudoverticillate or alternate, apically acute to rounded, 
often mucronulate, basally symmetric or asymmetric, acute, obtuse, or ta- 

PiPOLY, Cybianthus in Ecuador and Peru 67 

pering, rarely auriculate, the venation camptodromous, to brochidodromous, 
apically acute or attenute to rarely rounded, mucronulate or not, basally 
acute, cuneate, or rarely obtuse, at times subauriculate, symmetric or asymmetric, 
black or pellucid punctate, bearing hydropoten, at times with orange glan- 
dular lepidote scales or glandular papillae, the margin opaque or scarious, 
entire, subentire, undulate and sparse dentate or pectinate-dentate, flat or 
inroUed, the teeth vascularized or not; petioles pulvinate (basally swollen), 
canaliculate, except rarely marginate distally. Infioreicence pinnately to tripinnately 
paniculate, (rarely reduced to a pseudoraceme), pyramidal to columnar, the 
branches spicate or racemose (rarely glomerulate); inflorescence bract lin- 
ear-subulate, indistinguishable from the cataphylls; branch bracts membranous, 
linear, subulate; floral bracts linear-lanceolate, subulate, subtending or rarely 
on the pedicel, variously punctate, glandular-papillate or glabrous; pedicels 
terete, thin, or obsolete. Flowers unisexual or bisexual, 4-5(-6)-merous, 
homomerous or heteromerous; calyx cotyliform, at times unequally divided, 
the lobes valvate, spreading, linear-subulate or rarely subdeltate, apically 
acute, attenuate or rarely premorse, pellucid to black punctate, the margin 
entire or erose-serrulate, glabrous or glandular-ciliate; corolla rotate to subrotate, 
the lobes valvate, linear-subulate, apically acute, rounded or attenuate, es- 
sentially glabrous without, densely glandular-granulose throughout within, 
pellucid or black punctate, the margin irregular, entire; stamens and staminodes 
similar, the staminodes reduced in size, partially connate to form a con- 
spicuous or inconspicuous tube, at times bearing lobes alternating with apically 
free filaments, the tube adnate to the corolla, at times so as to mimic epipetalous 
stamens, the apically free portions of the filaments terete or flat, glabrous 
or glandular-granulose, the anthers dorsifixed, as wide or wider than long, 
apically rounded, obtuse or emarginate, basally cordate to deeply cordate, 
dehiscent by wide longitudinal slites; pistillode lageniform, clavate or conic, 
hollow, or entirely absent; pistil obpyriform, subglobose, obturbinate or clavate, 
the ovary terete, sparsely to densely translucent glandular lepidote, glan- 
dular-papillate or glabrous, the placenta umbonate, 2-3(-4)-ovulate, the 
ovules immersed in the placenta ca. 1/2 their length, the style short or ves- 
tigial, the stigma large, capitate, the margin lacinate, the lacinae with large 
lobes, the stigma early caducous. Fruit globose to depressed-globose, the 
exocarp sometimes fleshy, the mesocarp and endocarp stony, black or pellu- 
cid punctate, one-seeded, the testa corrugate, the embryo cylindrical, transverse. 

Distribution. — From the Darien of Panama southward through the Andes 
of Colombia, Ecuador, Peru and Bolivia to Rondonia, Brazil, at elevations 
of sea level-2,2()() m elevation. 

Ecology. — Members o^ Cybianthus subgenus Comomyrsim occupy the for- 
est floor, growing in sheltered areas under cliffs, and in other shady areas. 
The monoaxial stems are weak and break easily when trampled. They are 

68 Si DA 18(1) 

extremely sensitive to soil compaction and occur mostly in areas where leaf 
litter accumulates. 

Pipoly ( 1 987) demonstrated that Cyhianthus subgenus Comomyrsine is most 
closely related to subgenus Triadophora by the monoaxial habit. The cataphylls 
of Comomyrsme without petiolar structures, as opposed to the petiolate 
pseudocataphylls found in subgenus Triadophora, serve to further distinguish 
Comomyrsine. Likewise, the caducous, oversized stigma with lacinate lobes 
is unique to subgenus Commnynine. The subgenus contains 8 species, of which 
7 occur in Ecuador and Peru. 


1. Leaves 5.7-29 cm long; stems less than 1 cm diameter. 

2. Stems apices bearing orange hydropotes; leaf blades linear-lanceolate, 1.6- 
4.0 cm wide, the margin repand to undulate; petioles 0.6-1.0 cm long; 

inflorescence pseudoracemose, the flowers glomerulate 15. C. verticilloides 

2. Stem apices bearing rufotis glandular-papillate or with rufous hydropotes; 
leaf- blades ovate, obovate, oblanceolate, elliptic, or oblong, (4.0-), 10- 
22(-30) cm wide, the margin regular, entire or minutely straight-serru- 
late; petioles (I.O-)1.5-5(-21) cm long; inflorescence pyramidal-bipin- 
nate paniculate, or a columnar thyrsoid panicle, never a pscLidoraceme, 
the flowers spicate, corymbose, or rarely cymose. 

3. Stem apices sparsely glandular-papillate, bearing rtifous hydropotes; leaf 
bases broadly rounded, the margins scarious; petioles progressively longer 
acropetally along the stem, 2.0-1.5 cm long; inflorescence paniculate, 
the flowers corymbose; pethcels 1.0-2.5 mm long; sepals deltate; pet- 
als CLicullate apical ly, the margin glandular-granulose; filaments 

^1'iIt"()us 1(S. c. croatii 

5. Stem apices densely glandular papillate, without hydropotes; leaf bases 
acute, to acute with a rounded base, the margins opacjue; petioles subec]ual 
along stem, slightly shorter acroi^etally, 1.6-2.0 cm long; inflorescence 
pinnately paniculate, the flowers spicate or racemose; |uxlicels absent; 
sepals ovate-lanceolate; petals flat, the margin glabrous; filaments glandular 

MHinulose 17. c. humilis 

1. Leaves (16-)31-105 cm long; stems 1-5 cm diameter. 

4. Leaves oblong to elliptic, apically rounded or rarely acute, not mucrontilate, 
basally abruptly acute to broadly rounded; petioles ( (2.5-)5 cm long; flowers 
homomeroiis, 4— or 5-merous. 

5. Shrub or tree to 5 m tall; inflorescence a pyramidal |ianicle, the branches 
spicate, the flowers sessile to subsessile; leaf base symmetric; cataphylls 
(2.5-) 3.5-5.0 cm long 18. C. sprucei 

5. Semi-woody shnfl^is to l(-3.5)m tall; inflorescence a columnar, thyrsoid 
panicle, the branches racemose, the flowers on pedicels 2.3-5.0 mm 

long; leaf base asymmetric; cataphylls 0.6-2.5 cm long 19. C. simplex 

4. Leaves oblanceolate, apically acute, mucronulate, basally gradually tapered 
on the petiole, olten subauriculate; petioles 1 .5-2 cm long; flowers het- 
eromerous, tlie calyx 5-6-lobed, the corolla 4-5-lobed. 

6. Leaves chartaceous, 31-55 cm long, the margin flat, entire; cataphylls 

PlPOLY, Cybianthus in Ecuador and Peru 69 

1 .5-4.0(-4.5) cm long; calyx lobes deltate, the margin glabrous; co- 
rolla carnose 20. C. kayapii 

6. Leaves coriaceous, longer than 5 5-1 25cm, rhe margin mroUed, densely 
and minutely serrulate; cataphylls 4.0-6.5 cm long; calyx lobes linear- 
subulate, the margin densely glandular-granulose; corolla membrana- 
ceous 21. C. anthuriophyllus 

15. Cybianthus verticilloides (Cuatrec.) G. Agostini (Fig. 4B), Acta Biol. 

Venez. 10:165. 1980. Weige/fia vertni/Iozdes Cuatrec, Revisvj. Acad. Colomb. C'\. 
Exact. 8(31):327. 1951. Type: COLOMBIA. Valle del Cauca: Rio Call riverbed, 
between Quebradade Juntas and El Recreo, 2,070-2,260 m, 7 Jul 1946 (stam. R),J. 
Cucitrecasas 2 1981 (holcjtype: F; isotype: COL). 

Shrub to 1 m tall. Stem terete, 5-8 mm diam., sparingly rufous glandu- 
lar-papillate and bearing orange hydropotes, early glabrescent, the bark 
horizontally checked. Cataphylls chartaceous, subulate, (1.2— )1. 5-2.1 mm 
long, 0.9-2. 1 mm wide, flat, apically long-attenuate, basally somewhat decurrent, 
sparingly glandular-papillate and orange lepidote above and below, black 
punctate. Leaves pseud overticiUate; blades chartaceous, linear-lanceolate, 12- 
20 cm long, 1.6-4.0 cm wide, apically long-attenuate, basally acute, grey- 
ish-green above and pallid green below when fresh (teste Cuatrec), spar- 
ingly rufous glandular-papillate and orange lepidote above and below at 
first, persistent only below, inconspicuously pellucid-punctate, midrib planar 
above, prominent below, the secondary veins 8-12 pairs, prominent below, 
the margin inroUed except revolute basally, undulate to a vascularized blunt 
tooth at nerve end, translucent throughout its length but not scarious; petioles 
canalicualte, 0.6-1.0 cm long, 0.5-1.0 mm diam., glabrous, swollen ba- 
sally at point of attachment. Staminate Inflorescence: a pseudoraceme, 10—12 
cm long; peduncle ca. 1 cm long, the rachis glandular-papillate, the flowers 
glomerulate; inflorescence branch bracts membranous, subulate, 5.5—7.0 
mm long, 1.2-2.0 mm wide, minutely glandular-papillose publerulent, 
glabrescent, prominently black lineate-punctate, the margin entire; floral 
bracts similar to branch bracts except 0.9—2.0 mm long, 0.2—0.5 mm wide; 
pedicels obsolete to cylindrical to 1.3 mm long, prominently black punc- 
tate. Staminate flowers 5— 6-merous, lilac; calyx membranaceous, subcotyliform, 
1.5-2.3 mm long, unequally divided, the tube to 0.2 mm long, the lobes 
linear-lanceolate, subulate, 1.3-2.0 mm long, 0.4-0.7 mm wide, apically 
long-attenuate, flat, epunctate, glabrous, the margin irregular, entire, gla- 
brous; corolla rotate, chartaceous, 2.5-4.0 mm long, the tube opaque, 0.8- 
1.5 mm long, the lobes hyaline, ovate, 2.1-3.0 mm long, 1.4-1.9 mm wide, 
reflexed 180°, apically acute to rounded, moderately glandular-granulose 
throughout within except densely so along margin, glabrous without, epunctate, 
the margin irregular, entire; stamens 3.0-3.5 mm long, the tube 0.8-1.5 
mm long,epunctate, elobate, glabrous, the apically free portions of the fila- 

70 SiDA 18(1) 

menrs chick (base of filament as wide as anther), terete, glabrous, 1.5-2.0 
mm long, ventrally reHexed apically, the anthers subglobose, wider than 
long, 0.3-0.5 mm long and wide, apically rounded to obtuse, basally barely 
cordulate, the connective prominately black punctate dorsally; pistillode 
ovoid, ca. 0.9-1. .3 mm long, 0.6-0.8 mm diam. P ntillate infloresctna resem- 
bling staminate, but 4.5-6.5 cm long, the glomerules mostly reduced or a 
single flower; branch bracts and floral bracts identical, membranous, sublate, 
2.8-4.0 mm long, 0.7-1.0 mm wide, sparsely glandular-papillate, promi- 
nently black punctate, the margin erose; pedicels (1.5-)1.8-3.2 mm long. 
Pistillate flowers as in staminate but white; calyx chartaceous, cocyliform, 
2.2-2.7 mm long, hyaline, the tube 0.3-0.4 mm long, the lobes 1.9-2.3 
mm long, 0.7-1.1 mm wide, apically attenuate, at times rufous-papillate 
apically; corolla rotate, 2.5-3.2 mm long, the tube 0.9-1 .0 mm long, the 
lobes widely ovate, 1.6-2.2 mm long, 1.2-1 .5 mm wide, apically rounded 
to acutish, glandular-granulose within and along margin, the margin regular; 
staminodes similar to stamens but 1.6-1.9 mm long, the tube 0.9—1.0 mm 
long, the apically free portions of the filaments terete, 0.7-0.9 mm long, 
the antherodes globose, often malformed, 0.2-0.3 mm long, 0.3-0.4 mm 
wide, apically obtuse, basally obtuse to rounded or cordulate, the connec- 
tive prominently black punctate ventrally; pistil obpyriform, 2. 1-2.5 mm 
long, 1.2-1 .6 mm diam, the ovary 1.6-1.8 mm long, translucent-lepidote, 
glandular-papillate, prominently black punctate, style 0.5-0.7 nim long, 
the stigma capitate, 4-5-lobed, each lobe lacinate, to 0.6 mm long, the placenta 
deeply cupuliform, 2 ovulate. Pr/zit globose, 7-7.5 mm long and diam., 
prominently black punctate, exocarp thin. 

Distribution. — Previously considered endemic to the Western Cordillera 
of the Andes, on the cliffs (Farallones) of Call, in the Department of Valle, 
Colombia, at 1,900-2,600 m, but reported for the first time here from Napo, 
Ecuador, at 200 m elevation. 

Ecology and consen'ation status. — Cybianthus verticilloides is a rare species in 
lowland and premontane forest, growing near watercourses. Given that it 
has well-known uses, C. verticilloides might be suitable for cultivation. However, 
at present nothing is known regarding its natural population dynamics. Given 
the rapid urban development around Call, and its apparent rarity in Ecua- 
dor, this species should be considered endangered. 

Etymology. — The specific epithet refers to the strikingly pseudoverticillate 

Local names and uses. — Colombia: "verticilado" (Spanish); Ecuador: 
"Carnerupachapanga," "Yanacarneru" (Quichua), "Carnero Negro" (Spanish). 
Used to get rid of small fish in the urinary tract chat cause bleeding and pain. 

Specimens examineti. COLOMBIA, Valle del Cauca: Ri'o C^ali rivcrbetl, above Ri'o Pichinde, 
El Rnbal, IM^") m, 25 Jul 1 946 (fr),_/, Ciuitmciscis 2 1 12 1 (COL, F); Penas Blancas, 2,200 

PiPOi.Y, Cybianthus in Ecuador and Peru 71 

m, 9 May 1940 (stam. fl), A. Figueroa 873 (COL, US); Rfo Tulua, 1,200 in, without date 
(pist. fl),./. Duque-Jaramilk 4022 (COL); El Silcncio, Yanaconas, 1,900-2,200 m, 28 Feb 
1939 (pist. fl), E. Killip & H. Garcia 33802 (COL, US); Farallones de Cali, 1,800 m, Oct 
1883 (pist. fl), F. Lehmann 3027 (K, US); km 18-20, Cali-Buenaventura Hwy, entering 
near Finca Zingara, summit of the Cordillera Occidental, 1,500-2,000 m, 28 Feb 1988 
(fr), H. van derWerff& I. Cabrera 15786 (COL, MO, VALLE). ECUADOR. Napo: Rfo 
Napo, S bank a few km below Itaya, 00° 28' S, 76° 33' W, 200 m, 20 Aug 1982 (ster.) H. 
Balslev & Santas Dea 2850 (QCA). 

Cybianthus vertkilloides appears to be most closely related to C. goudotianus, 
by the synapomorphic vascularized leaf teeth. However, C. verticilloides is 
defined by the autapomorphic horizontally checked bark, the orange hydropotes 
of the branchlets, and the linear-lanceolate leaf shape. 

16. Cybianthus croatii Pipoly, sp. nov. (Fig. 4A, 16). Type: ECUADOR. Pastaza: 
Along Rd. between Diez dc Agosto and Arajuno, 18 km NE of main Puyo-Macas 
Rd., 8.2 km NE of Diez de Agosto, 01° 27' S, 77° 51' W, 970 m, 4 May 1984 (stam. 
fl), r. Croat 59009 (hoi.otyfp: MO). 

Quoad habitum deminutum petioles brevistipitatos et laminas chartaceas, C. hiimilem 
valde cognatum, sed ab ea basibus laminaribus obtusis vel rotundatis (non acutis) inflorescentiis 
paniculatis cum ramulis floriferis corymbosis (nee tripinnati- paniculatis cum ramulis floriferis 
spicatis vel racemosis) petalis ad apicem cucullatis (nee planis) ad marginem glandulari- 
papillosis (nee glabris) filamentis glabris (nee glandulari-granulosis) antherarum connectivis 
manifeste punctatis (nee epunctatis) prompte cognoscitur. 

Suhshrub to ca. 15 cm tall. Stem terete, ca. 4 mm diam., bearing rufous 
hydropotes and sparingly glandular-papillate. L&ives alternate; blades chartaceous, 
ovate to elliptic, 5.7-14 cm long, 4.0-7.5 cm wide, apically acute, the tip 
mucronulate, basally obtuse to rounded slightly decurrent on the petiole, 
bearing rufous hydroporen above and below, midrib slightly impressed distally, 
slightly raised proximally above, prominent below, the secondary veins 4- 
9 pairs, barely visible above, prominent below, without collecting vein, the 
margin scarious, subentire or entire; petiole deeply canaliculate, decurrent 
on stem, at times appearing to form a small sheath, (1.5-)2.0-4.5 cm long, 
0.2-0.3 cm diam., increasing in length acropetally along stem. Cataphylls 
membranaceous, alternate, subulate, 6-12 mm long, 0.6-1.2 mm wide, 
located just below center of internode, psuedoverticillate, apically acicular, 
keeled, conspicuously black punctate-lineate, bearing hydroporen and glandular 
papillae. Inflorescence bract subulate, 3.9-4.0 mm long, 2.0-3.0 mm wide. 
Staminate inflorescence: supraaxillary, paniculate, 1.2-3.5 cm long, appearing 
succulent, the branches racemose, the rachis densely glandular-papillate; 
peduncle 0.5-1.0 cm long; inflorescence branch bracts linear-lanceolate, 1.8- 
2.2 mm long, 0.4-0.6 mm wide, conspicuously punctate, glandular-papil- 
late; floral bracts subtending and equal to the pedicels, 1.0-2.5 mm long. 
Staminate jhwers homomerous, 4-merous, green; calyx cotyliform, membranaceous, 
1.2-1.5 mm long, more or less equally divided, hyaline, the tube 0.2-0.3 


SiDA 18(1) 

Fig. 16. Cybianthiis crvattt Pipoly. A. Habit, showing stem with small, acicular cataphylls, 
atropetally longet petioles, and supraaxillary inflorescences. B. Staminate flower bud, showing 
sparsely glandular-papillate, cotyliform calyx. C. Open staminate corolla, showing cucul- 
late lobe apices, suborbicular anthers and conic pistillode. D. Open pistillate corolla, showing 
oblate antherodes, subglobose pistil. A-C, drawn from holotype. D, drawn from L. Albert 
ck Escobar il 44- Figtire drawn by Peggy Duke. 

PiPOLY, Cybianchus in Ecuador and Peru 73 

mm long, the lobes deltate, 1.0—1.2 mm long and wide, apically acute to 
somewhat acuminate, prominately black punctate, sparsely glandular-papillate 
without, the margin irregular, entire, glandular-papillate at first, glabres- 
cent; corolla subrotate, carnose, 2.4—2.7 mm long, the tube 0.7—0.8 mm 
long, translucent, the lobes opaque, oblong, 1 .7—2.1 mm long, 0.9— l.O mm 
wide, apically rounded to obtuse, cucullate, sparingly glandular-papillate 
without, densely glandular-granulose within, prominently black punctate 
without, especially at apex, the margin entire, glandular-granulose; stamens 
2.5—2.6 mm long, the tube conspcuous, carnose, 0.7—0.8 mm long, subtruncate, 
the apically free portions of the filaments terete, 0.9-1-0 mm long, epunctate, 
glabrous, erect except slightly recurved ventrally at point of attachment to 
anther, the anthers suborbicular, 0.7—0.8 mm long and wide, apically rounded, 
deeply cordate basally, the connective epunctate ventrally, prominently black 
punctate dorsally; pistillocle conic, 1.3 mm long, 0.7 mm wide, densely 
translucent-lepidote, conspicuously black punctate, hollow. Pistillate inflorescence: 
as staminate, but 2.5—3.5 cm long; pedicels 0.6—1.1 mm long. Pistillate 
flowers as in staminate but calyx 1 .3-1-5 mm long, equally divided, the tube 
0.2—0.3 mm long, the lobes triangular, 0.9— 1-2 mm long, 0.5—0.7 mm wide, 
apically acuminate, sparsely glandular-papillate along margin without; corolla 
subcampanulate, the lobes 1.3-1-5 mm long, the tube to 0.2 mm long, the 
lobes suborbicular, 1.1—1.3 mm long, 1.0—1.1 mm wide, apically rounded 
to obtuse, minutely glandular-granulose along margin without and throughout; 
staminodes as in stamens but 0.9—1-0 mm long, the tube, ca. 0.2 mm long, 
the apically free portions of the filaments 0.2—0.3 mm long, the antherodes 
oblate, 0.4—0.5 mm long, 0.5—0.6 mm wide, apically truncate, basally obtuse, 
the connective prominently black punctate ventrally; pistil subglobosel, 0.6— 
0.7 mm long and diam., densely translucent-lepidote, conspicuously black 
punctate, the placenta deeply cupuliform, bearing 2 ovules, the style barely 
discernible, to 0.1 mm long, stignma subcapitate, the margin lacinate, early 
caducous. Fruit unknown. 

Distribution. — Known only from the type (Ecuador) and from Antioquia, 
Colombia, at 970—2,440 m elevation.. 

Ecology and conservation status. — Cybiantbus croatii occurs in premontane 
and montane wet forest, where it is locally common in protected areas near 
forest margins. Because of its restricted habitat, it is presumed to be threatened. 

Etymology. — It is with great pleasure that 1 dedicate this species to Tho- 
mas B. Croat, pre-eminent authority on the systematics of Neottopical Araceae, 
indefatigable collector, gentleman, scholar, and the P. A. Schuize Curator 
of Botany at the Missouri Botanical Garden. During the tropical botany 
course I took from him in Costa Rica in 1977, 1 was first shown and be- 
came intrigued with the systematics and populaton biology of the Myrsinaceae. 

74 SiDA 1<S(1) 

ParA'i YFi:: COLOMBIA. Antioquia: Mpio. Caklas, above town, Finca La 7:drz.d, 2, 140 
m, 2 Feb 1984 (pist. H), L. Albert cle EsaAir et al. 3744 (HLJA, US). 

Cyhiaiitb//s crocit'a is the smallest myrsinaceous shrub chat exhibits Corner's 
Model (Halle et al. 1978). It is interesting to note that with each successive 
pseudowhorl of cataphylls, an inflorescence is produced, followed by a larger 
leaf with a longer petiole. This species appears to be most closely related to 
another diminutive plant, C. humilh, known only from the Department of 
Antiocjuia, Colombia and adjacent Choco, and from Ecuador. The synapomorphy 
which defines the Cybtanthus humllis-C. croatii clade is the scarious leaf margin, 
known elsewhere only in subgenus Grammadenia (Pipoly 1 987, unpubl. data). 
However, becatise that subgenus occurs on the other side of the generic cla- 
dogram presented by Pipoly (1987), it is most parsimonius to hypothesize 
that the margin has arisen independently in these distant lineages. The pistillode 
is present in both of these species, and I have chosen to postulate that this 
represents a reversal because it is the most parsimonius conclusion. Cybianthns 
croatii is defined by the autapomorphic rufous stem hydropotes and the ac- 
ropetally longer petioles. Despite the fact that the distribution of this spe- 
cies entirely overlaps that of C, hnniilis, it appears that C. croatii is restricted 
to montane and cloud forests, where C. h/nnilis is restricted to premontane 
pluvial forests and subparamo thickets. This is yet another example of sis- 
ter species in alritudinally adjacent liabitats, already reported in Cybia)!tb//s 
subgenus Laxiflor//s (Pipoly 1983) C. subgenus /\i/rmY;;/'r;wr;r/7/:^^ (Pipoly 1983b) 
and C. subgenus Conormrpha (Pipoly 1992a). Whether this supports the conce[it 
of speciation by peripheral isolation (parapatric) is unknown. Futher stud- 
ies of the respective population biologies of the subgenus are needed. 

17. Cybianthus humilis (Mez ) G. Agostini (Fig. 3F), Acta. Biol. Venez. 
10:163. 1980. \Veigelthih/(mil,sUi:z in EngL, Pflanzenr. IV. 236(Hetr 9):291 . 1902. 
Typh: ECUADOR. Without locaHty, 1896 (stam. fl), A. Sodiro 100/14 (holotyph: 
B-destr., F Neg. 4856;'icnYPE, here designated: COLOMBIA. Antioquia: 2,650- 
2,800 m, 1 Apr 1880 (stam, H), W: Kdlhreyer n34 (K). Altliotigh Mez (1902) did 
not specifically mention the Sodiro collection as the t\pe, he cited the Sodiro and 
the Kalbreyer specimens in the protologtie. However, the F photograph clearly shows 
that a drawing of a dissection accompanies the Sodiro specimen, while that of Kalbreyer 
does not, stiggesting that the Sodiro specimen (ormed the |irincipal element upon 
which the description was based. LJnfortimately, no duplicates of that Sodiro collecton 
have been located. Mez also annotated the Kalbreyer sheet at K, and thus I designate 
it as the lectotyjie. 

CoiitoD/yr.iiiii' soclirihnni Mez, Btill. Fierb. f^oissier, 2 ser 5:535. 1905. syn. nov. Cyb'uinthin 
mimkiniis (Mez) C. Agostini, Acta Biol. Venez. 1 0: 1 63. 1 980. Tyi'p: ECUADOR. Atacat/o. 
Jan 1902 (stam. fi), A. Sodivn 100/2 (iioi.otypf.: B-destr.; i.H;r()rYPP., here designated: P). 

Sabshri/b to 25 cm tall. StcDi terete, 3.5-4.0 mm diam., densely glandu- 
lar-papillate, early glabrescent. Cataphylls in alternating nodes with leaves, 
membranaceous, subulate, 1 0-13 mm long, 1 .7-2.6 mm wide, apically long- 

PiPOLY, CybianchLis in Ecuador and Peru 75 

attenuate, keeled, midrib prominent below, prominently black punctate and 
lineate-ptmctate, glandular-papillate, glabrescent, the margin opaque, flat, 
entire. Leaves subopposite; blades chartaceous, elliptic to narrowly elliptic, 
1 5— 20(— 27) mm long, (4.0— )7. 0—8. 5 cm wide, apically acute to subacuminate, 
terminating in an inconspicuous mucro, basally acute, bearing hydropoten 
above and below, inconspicuously pellucid punctate, midrib somewhat impressed 
above, prominent below, the secondary veins 6—1 1 pairs, slightly impressed 
abive or not, prominently raised below, not united by a submarginal col- 
lecting vein, the margin entire, opaque, subrevolute; petioles canaliculate, 
thin, 1.0—2.0 cm long, 2.0—2.5 mm diam., glabrous, not decurrent on the 
stem. Infiorescence bracts similar to cataphylls, but 1.2—1.3 mm long, 1.5— 
2.2 mm wide. Staminate inflorescence: a bipinnate panicle, 3-5— 15(— 19) cm 
long, the rachis densely glandular-papillate, the branches spicate or rarely 
racemose, appearing subglomerulate apically; inflorescence branch bracts 
membranaceous, subulate, 3—4.5 mm long, 0.3—0.5 mm wide, densely and 
prominently black pimctate-lineate, sparingly papillate; pedicels essentially 
obsolete, to 0.3(— 2) mm long; floral bract membranaceous, ovate, asym- 
metric, 2.3—2.8 mm long, 0.9-1 -2 mm wide, apically abruptly acuminate, 
medially keeled, epunctate, the margin erose, stipitate glandular-papillate. 
Staminate flowers 4— 5-merous; calyx membranaceous, cotyliform, 1.5—2.0 mm 
long, tube 0.2—0.3 mm long, the lobes ovate to lanceolate, 1.3—1.7 mm 
long, 0.5—1.3 mm wide, unequally divided, apically acuminate to acute and 
often erose or premorse, glandular papillose-puberulent without, the mar- 
gin erose, glabrous; corolla chartaceous, appearing subrotate, 3-7-4.5 mm 
long, the tube 1.1—1.5 mm long, the lobes linear-lanceolate to oblong, 2.4— 
3.2 mm long, 1.0—1.5 mm wide, often unequal, apically long-attenuate to 
obtuse, moderately rufous gladular-granulose throughout within, sparsely 
glandular-papillate without, hyaline, the margin entire, glabrous; stamens 
3-0-3.9 nnm long, the tube 1.1—1 .5 mm long, coriaceous, conspicuous, sparsely 
glandular-grandulose, elobate, the apically free portions of the filaments basally 
as wide or wider than anther, then tapering apically, 1 .9— 2.4 mm long, terete, 
glandular-granulose, the anthers oblate, 0.4—0.5 mm long, 0.6—0.7 mm wide, 
apically truncate, basally subcordate, the connective epunctate; pistillode 
glabrous lagenform, 1.4—1.6 mm long, 0.8-1.0 mm wide, the stigma punctiform. 
Pistillate inflorescence: as in the staminate but 7—10 cm long, 4—6 cm wide; 
inflorescence branch bracts 2.5-3.5 mm long, 0.2—0.3 mm wide; peduncle 
1—2.5 cm long; floral bracts 1 — 1.5 mm long; pedicels obsolete. Pistillate 
flowers unknown; fruiting calyx as in staminate but 1.3-1.5 mm long, the 
tube ca. 0.2—0.3 mm long, the lobes 1 .1 — 1 .2 mm long, 0.2—0.3 rnm wide. 
Fruit globose, reddish-purple when fresh, 4—6 mm long and diam. when 
dried, the exocarp thin, conspicuously pellucid punctate. 

76 SiDA 18(1) 

Distribution. — Known only from Colombia and Ecuador, at 1,450—2,700 
m elevation. 

Ecology and conservation status. — Cybianthus hinnilis grows in deep shade 
among rocks at the margins of premontane pluvial forests. Populations I 
observed in Colombia grew only in undisturbed areas with deep shade, leaf 
litter and organic humus. Because of the apparently restricted habitat, this 
species should be considered threatened. 

Etymology. — The specific epithet refers to the low habit of the plant. 

Local names and uses. — Colombia (Choco): "Hierba del palo grande." Ground 
to make crude syrups; syrup applied externally to cure cuts, taken inter- 
nally to cure internal infections, clean the stomach and against chills. Given 
that it is a shade species, the common name probably refers to its frequency 
in sliaded areas near large trees. 

Si^cci mens examined. COLOMBIA. Antioquia: Mjiio. Urrao, i-'arque Nacional Uis Orqui'deas, 
VeredaCalles, permanent inventory tor premontane pluvial iorest, right bankof Ri'o Calles, 
06° 32' N, 76" 19' W, 1,450 m, 27 Nov 1993 (fr),./. P,poly et at. 17186 (COL, JAUM, 
MO), 1,450-1,500 m, 29 Nov 1993 (lr),_/. Pi/w/) via/. 17361 (COL, JAUM, MO); with- 
out locality and date, (.stam. fl), F. Lehiiiatni s.u. (L, K). Choco: Mpio. de Quibdo, C^orregimiento 
San Francisco Icho, Quebrada Caledonia along Caledonia Rd., 9 Apr I 9H7 (ster.), F. Garcni 
& J. Ecl:uirary/d 259-A. (COL, CHOCO, MO). Nariiio: Mpio. Barbacoas, Corrcgimiento 
Orri'z y Zamora, Vereda El Barro, Reserva Natural Rfo Nambf, ca. 5 km W de Altaquer, 
taldas occidentales de la Cordillera Occidental, 01" 18' N, 78° 08' W, 1,350-1,100 m, 3 
Sep 1997 (ster.),/ P//W). A. Cngollo. et ul. 21240 (BRIT, rMB,JAUM, PSO). Quindio: 
Mpio. De Salento, Estacion Navarco, Alto San Ignacio, 2,850 m, 23 Nov 1990 (stam. fl), 
P. Franco et ul. 3204 (COL, MO). Risaralda: Mpio. Sta. Rosa, Camino de LIcrradura etre 
Termales y Paramo Sta. Rosa, Cordillera Central, vertiente Occidental, Hacienda El Margariral, 
2,500 m, 18 Atig 1980 (fr),/ hlrolmetal. 967/ (COL, MO). ECUADOR. Napo: Canton 
El Chaco, Proyecto Hidroelectrico Coca, Punto ST3, right bank of Ri'o Quijos, ca. 10 km 
S of Reventador, 1,500 m, 3-5 Oct 1990 (fr), W. Pjlacim 5930 (MO, QCNE). Pichincha: 
Cant()n Quito; Parroquia Calacah', Reserva Geof-)otanica Pukiiahua, 00° 01' N, 78° 35' W, 
1,800-2,000 m, 29 Jul I 989 (stam. fl), C. Cerm 7 1 H4 (MO, QCNA); Mindo, 26 Jun 1876 
(stam. fl), £. Andre 3819 (K). 

Cybianthus bumilis is most closely related to C. croatii, by virtue of its 
synapomorphic scarious leaf margin. The autapomorphies that distinguish 
C. humilis from all other species of the subgenus include the premorse apices 
of the calyx lobes, the unecjual corolla division, and the oblate anther shape. 

18. Cybianthus sprucei (Hook, f.) Cj. Agostini, Acta Biol. Venez. 10(2): 164. 

1980. Couioniyrsine iprncei Hook, I. ni Benth. et Hook., Cien. PI . 2:614. 1876. Weigeltia 
spriicei (Hook, f.) Mez in Engl., Pflanzenr. IV. 236(Heft 9):291 . I 902. Type: ECUA- 
DOR. [CiiiMBORA/.o: W slopes of Volcan Chimborazo, 17 Jun 1860] (stam. fl), 11. 
Spruce 6)144 (houxitpe, K-2 sheets). Note: label on specimen does not indicate place 
or date. Information was derived by comments regarding habit, etc. on the label which 
matched data gi\'en in Sprtice ( I 880). 

Weigeltia pitncimensis Standi., Publ. Field Mus. Nat. Hist, Bot. Set. 22:l6'l. 1940. syn. 
nov. Cyhicnitljus paiuiinensis (Standi.) CL Agostini, Acta Biol. Venez. 1 0: 163- 1980. Type: 

PiFOLY, Cybianthus in Ecuador and Peru 77 

PANAMA. Darien: Cana, Cuasi Trail, Dtto. Cheijana, 1000 m, 10 Mar 1940 (stam. 
fl), M E. Terry & R. A. Terry 1490 (holotype: F-2 sheets; isotypes: A, MO). 
Weigeltia purpurea Cuatrec, Revista Acad. Colomb. Ci. Exact. 8(31):326. 1951. Type: 
COLOMBIA. Valle del cauca: Bahia de Buenaventura, Quebrada de San Joaqui'n, 
0-10 m, 21 Feb 1946 (stam. fl),J. Cuatrecasas 19892 (holotype: F; isotype: COL). 

Shrub to tree to 5 m, flowering from less than 1 m. Stem terete, 1.3-2.5 
cm diam. below uppermost leaves, swollen at nodes, semi-woody, glandu- 
lar-papillate-pLiberulent, glabrescent. Cataphylls few, alternate, coriaceous, 
subulate, (2.5-)3.0-5.0 cm long, 2-4 mm wide, strongly keeled, densely 
glandular-papillose-puberulent, glabrescent, consiptiously black punctate, 
the margin opaque, regular, entire. Letft'(?j-pseLidoverticillate; blades chartaceous, 
widely oblong to elliptic, rarely widely obovate, (26— )3 1—7 5 cm long, (6.5—) 
10—30 cm wide, apically rounded, obttise or rarely acutish, not mucronulare, 
basally abruptly subcuneate, asymetric, slightly decurrent on petiole, bear- 
ing a few hydropoten above, moderately rufous glandular-papillate and with 
a moderate number of hydropoten below, prominently red or black puctate, 
the margin regular, opaque, entire; petioles stiff, (2.5-)5.0-l4(-23) cm long, 
0.3-0.9 cm diam., slightly to moderately canaliculate, swollen basally, sparingly 
glandular-papillate, glabrescent. Staminate Inflorescence: a pyramidal bipin- 
natethyrsoid panicle, 9-l6.5(-19)cm long, 9-17(-22)cmwide, the branches 
subspicate, the rachis densely glandular-papillose-puberulent; peduncle 0.5- 
2.0 cm long; inflorescence branch bracts chartaceotis, linear-lanceolate, 8.5— 
12.9 mm long, 0.6—0.9 mm wide, apically narrowly acute, conspicuously 
black punctate, densely glandular-papillate, gabrescent, flat, the margin opaque, 
regular, entire; floral bracts membraneous, linear, 1.5— 2.0(-2.5) mm long, 
0.5—0.6 mm wide, subulate, hyaline, densely glandular-papillose puberu- 
lent, the margin entire, glandular-papillate; pedicels obsolete to 0.2 mm 
long. Staminate flouers homomerous, 4-merous, coriaceous, subscssile, race- 
mose, 4— 5-merous, white in btid, green in anthesis, then crimson; calyx 
coriaceous, subcotyliform, unequally divided, 0.9—1-1 mm long, the tube 
0.1—0.2 mm long, the lobes linear-lanceolate, 0.7—0.9 mm long, 0.2-0.5 
mm wide, apically subulate, keeled, brown punctate-lineate or punctate medially, 
sparsely glandular-papillate, the margin irregular, subentire to erose, densely 
glandular-ciliate; corolla subrotate, 2.0-2.9 mm long, the tube 0.2— 0.3mm 
long, the lobes narrowly ovate to lanceolate, (1.7— )1.9— 2.3(-2.6) mm long, 
0.9-l-0(-1.5) mm wide, reflexed 135° from tube at anthesis, apically at- 
tenuate, densely glandular-granulose throughout within and along margin 
within and without, inconspicuously pellucid punctate, the margin entire, 
somewhat irregular, densely glandular-granulose; stamens exserted to slightly 
shorter than the corolla lobe, 1.6—2.2 mm long, the tube ca. 0.5 mm long, 
conspicuous, coriaceous, taller than the corolla tube, elobate, opaque, epunctate, 
glabrous, the apically free portions of the filaments terete, thicker than the 

78 SiDA 18(1) 

anthers, 0.8-1.6 mm long, ventrally recurved at anthesis, epunctate, gla- 
brous, the anthers subglobose, ca. 0.3 mm long, 0.4-0.5 mm wide, apically 
rounded, basally cordulate, dorsifixed near base so as to appear basihxed, 
the connective prominently red or black punctate dorsally; pistiUode nor- 
mally absent, occasionally conic, to 1 mm long, 0.3 mm wide, densely translucent- 
lepidote. Pistillate hiflorescence resembling staminate in all features, but smaller, 
2. 5-3. 5(-l 3) cm long, 3.5-8. 0(-l 2.5) cm wide; peduncle 0.5-1 .5 cm long; 
inflorescence branch bracts 3.0-3.5(-5.0) mm long, to 0.6 mm wide, at 
times somewhat cucuUate; floral bracts 2.0-2.6 mm long, ca. 0.5 mm wide, 
pedicels virtually obsolete, or cylindrical to 0.1 mm long in flower, accrescent 
and incrassate to 2.0 mm long, 1.0-1 .5 mm diam. in fruit. Pistillate flowers 
as in staminate, forming a condensed spike on the inflorescence branches; 
calyx subcupuliform, 1.2—1.4 mm long; the tube to 0.1-0.2 mm long, the 
lobes 0.9-1.1 mm long, 0.9-1-0 mm wide, widely ovate, apically acute; 
corolla rotate, 2.7-2.9 mm long, the tube 0.6—0.7 mm long, the lobes ob- 
long to elliptic, 2.0—2.2 mm long, 0.9—1 .0 mm wide, apically acute; staminodes 
1.5—1 .6 mm long, the tube 1 .0—1. 1 mm long, the apically free portions of 
the filaments 0.4-0.5 mm long, the antherodes malformed, 0.2-0.3 mm 
long, 0.3—0.4 mm wide; pistil obturbinate, 1.5-1 .6 mm long, densely trans- 
lucent-lepidote and prominently pellucid punctate, the style obsolete, the 
stigma capitate, 0.2—0.3 mm long, subsessile, 4— many-lobed, the lobes 1.3— 
2.3 mm long, viscid, bright crimson, the placenta ovoid, the ovules 2, born 
on side of placenta. Fr//it subglobose, 5.0—9-0 mm long, 9-14 mm diam., 
fleshy, the exocarj:> thick, orange at maturity, prominently black punctate. 
Bisexual Inflorenscence resembling staminate in all features except: 3-7-14.5 
cm long, 2.5-12.5 cm wide; inflorescence branch bracts and floral bracts 
and pedicels as in pistillate. Bisexual flotuers spicate, less crowded than in 
the pistillate, more crowded than staminate; calyx cotyliform, (4-)5 -parted, 
1.0—1.2 mm long, the tube ca. 0.2 mm long, the lobes narrowly ovate to 
narrowly triangular, 0.8-1.0 mm long, 0.3—0.4 mm wide, apically acute 
to narrowly acute; corolla 4-lobed, rotate, 2.0—2.7 mm long, the tube ca. 
0.3 mm long, the lobes oblong, 1 .7—2.4 mm long, 0.9—1 -2 mm wide, apically 
acute; stamens 1.2-1.5 mm long, the tube ca. 0.5 mm long, the apically 
free portions of tlie filaments 0.6—0.7 mm long, the anthers oblate, 0.3— 
0.4 mm long, ca. 0.5 mm wide; pistil as in pistillate flowers except 1.2—1.4 
mm long, the ovary 1.0—1.1 mm long, 0.7—0.9 nim diam., the style short, 
0.1—0.2 mm long, the stigma capitate, to 0.2 mm long, the lobes 0.9-1-1 
mm long, the placenta ellipsoid, the ovules 2—3, borne on side of placenta. 
Bisexual fruit s\xh^\ohose, as in pistillate, but 4-5 mm long, 5-7 mm diam. 

Distribution. — From Darien, Panama to Loreto, Peru, from 0—1,700 m 

Ecology and conservatifm status. — Cybianthus sprucei is a ridgetop species. 

PiPOi.Y, Cybianrhus in Ecuador and Peru 79 

occurring in premontane wet and rainforests and also in lowland forests on 
forest margins of exposed hilltops. This species, as opposed to its closest 
congener, Cybianthus simplex, occurs in areas of high incident light for at 
least part of the day. Because Cybianthus sprucei continues to reproduce in 
spite of mild forest intervention, it is not considered threatened or endan- 

Etymology. — This species was named in honor of Richard Spruce, ardent 
collector and student of the Andean and Amazonian flora. 

Local names and uses. — Colombia: "margoandre," "Tunda" Spanish (Valle 
del Cauca, Colombia); Ecuador: "urcu tahucu" (Quichua); ground and a vapor 
bath is taken to "send evil away" and to treat body pains, headaches, inter- 
nal colds; Peru: "kurup" (Jfvaro); the root is mashed and boiled, and the 
decoction is drunk to "strengthen" the body. 

Specimens examined. PANAMA. Darien: Orro Pirre, 10-20 Jul 1977 (bisex. fl, it), J. 
PoLuwi 4544 (MO, PMA); on ridge of Cerro Pirre, 08° 00' N, 77° 45' W, 1,000-1,080 m, 

14 Sep 1989 (fr), G. McPherson 14066 (BRIT, MO). COLOMBIA. Cauca: Rfo Micay, en 
Guayabal, 5-20 m, 25 Feb 1943 (pisr. fr),/. Cua/recasas 1415B (COL, F, US); Distrito Cauca, 
El Tambo, 900 m, Apr 19.37 (stam. fl), K. imi Sneidern 1615 (S). Choco: Along Rd. be- 
tween Quibdoand Medellin, Km 207.5, 0-200 m, 18 Dec 1980 (pisr. fl, fr), T. Croat & A. 
Cogollo 522^7 (COL, JAUM, MO); Alto del Buey, 1,200-1,800 m, 8 Jan 1973 (pist. fl, fr), 
A. Gentry & E. Forero 73/7 (COL, F, MO). Narifio: La Guayacana, Funes, 24 Jun 1951 
(stam. fl), R. Castatleda 2873 (COL, F); Ricaurte, 1 ,300 m; 18 Apr 1941 (bisex. fl), K. von 
Sneidern A61 2 his (S); Reserva Natural La Planada, 7 km above Chucunes on Rd. between 
Tuquerres and Ricaurte, along Sendero La Vieja, 01° 06' N, 77° 54' W, 1,780-1850 m, 7 
Mar 1990 (fr), T. Croat 71155 (MO, PSO). Valle del Cauca: Pacific coast, Rfo Naya, Puerto 
Merizalde, 5-20 m, 22 Feb 1943 (bisex. fl),/. C/u/trecasas 14053 (COL, F); Rfo Yurumangui, 
550 m, 28 Jan-10 Feb 1944 (pist. fl, fr),J. C/tatrecasas 15743 (COL, F, US); Rio Calima, 
Choco region. La Trojita, 5-50 m, 20 Feb 1 944 (stam., pist. fl-sheets mixed),J. Cuatrecasas 
16272 (COL, F, US); Rfo Cajambre, Barco, 5-80 m, 30 Apr 1944 (fr),/ Cuatrecasas 1 7625 
(COL, F, US); Bajo Calima, 15 km N of Buenaventura, Carton de Colombia concession, 
Juanchacho region, 03° 56' N, 77° 08' W, 500 m, 27 Mar 1986 (stam. fl), A. Gentry et al. 
53713 (COL, MO), Concesion Pulpapel/Buenaventura, 03° 55' N, 77° 00' W, 100 m, 7 
Mar 1985 (fr), M. Monsa/ve 767 (COL, CUVC, MO), 1 9 Mar 1 985 (stam. fl), M. Monsalve 
790 (CUVC, MO), 21 Mar 1985 (sram. fl), M. Monsaive 797 (CUVC, MO), 12 Mar 1986 
(stam. bud), M. Monsalve 981 (CUVC, MO), 24 Aug IS'86 (fr), M. Monsaive 1 124 (CUVC, 
MO); Bahia de Malaga, near moutii of Quebrada La Sierpe,0 4° 00' N, 77° 1 5' W, 0-20 m, 
17 Feb 1983 (stam. fl), A. Gentry et aJ. 40453 (COL, MO); 1 8 km E of Buenaventura, 50 
m, 14 Feb 1939 (fr), E. Killip & H. Garcia 33219 (BM, COL, NY, US), Buenaventura, Jun 
1901 (pist. fl, fr) F. Lelomann B. T 651 (K, NY). Putumayo: Umbna, 00° 54' N, 76° 10' 
W, 325 m,Jan-Feb 1931 (stam. fl), G. Klug 2108 (US). ECUADOR. Bolivar : Along first 

15 km of ChiUanes-El Tambo, 2,400 m, 18Jul 1991 (stam. fl), H. van clerWerffet al. 12430 
(BRIT, MO, QCNE); along Rd. ChiUanes-San Pablo, 6 km outside Chillanes, 2,600 m, 21 
Jul 1991 (fr), H. vander Werffetal.l 12561 (MO, QCNE), Morona-Santiago: 15 km N of 
Macas, Rd to Rio Upano, 02° 07' S, 78 °08' W, 1250 m, 20 Feb 1987 (bisex. fl),^. Bohlin 
et al. 1493 (GB); Cordillera de Cutucu, W slopes along trail from Logrono to Yaupi, 02° 
46' S, 78° 06' W, 1,200 m, 10 Nov 1976 (stam. fl), Al. Maddtson et aL 3204 (US). Napo: 
Canton Archidona, Carretera Hollfn-Loreto, Rfo Huataraco, 2 hrs walk from Guagua Sumaco, 

80 SiDA 18(1) 

00° 43' S, 77° 32' W, 800-1,000 m, 23-30 Aug 1989 (stain, fl), C. Ceron & M. Factos 
7648 (MO, QCNE); Canton Orellana, Resetva Flori'stica El Chuncho, 5 km N of Coca, 00° 
25' S, 77° or W, 250 m, 23 May 1993 (fr), W. Palados 10680 (MO, QCNE), Ei Chuncho, 
el Payamino, Estacion Experimental INIAP-Napo, 5 km NW of Coca, 00° 30' S, 77° 01' 
W, 250 m, 12 Oct 1987 (stam. fl), C, Cerd/i 2494 (MO, QCNE); Canton Tena, Mishualli, 
junction of Rios Mishualli and Napo,01° 03' S, 77° H ' W, 500 m, 13-14 Aug 1979 (fr), 
L. Hohii-Niehoti 19295 (AAU); Estacion Biologica Jatun Sacha, along S bank of Ri'o Napo, 
1 km E of Puerto Misahualli, 00° 04' S, 77° 36' W, 450 m, 1 Apr 1992 (fr), T. Croat 73552 
(MO, QCNE), 8 km E of Puerto Mishualli, 01° 04' S, 77° 36' W, 400 m, 14 Aug 1989 
(ster.), C. Ceron 7409 (MO, QCNE), 22 Sep 1989 (stam. fl), W. Palacios 4471 (MO, QCNE), 
16 Mar 1991 (fr), D. Neill98J3 (BRIT, F, MO, QCNE); Ri'o Blanco Comunidad, headwa- 
ters of Rio Huambuno, 6 km NNW of Ahuano, 01° 44' S, 77° 44' W, 440 m, 18 Jul-9 
Aug 1990 (fr), E. Kohn 1311 (MO). PERU. Amazonas: Prov. Bagua, Yamayakat, trail to 
Putuim, 04° 55' S, 78° 19' W, 500 m, 17 Oct 1996 (stam. fl), R. Va'squez & N. Jarcmillo 
20318 (AMAZ, BRIT, MO). Loreto: Prov. Loreto, Pampa Hermosa and vicinity, Rio Corrientes, 
1 kmSofjunctionwithRjoMacusari,03° 15' S, 75" 50' W, l60m, 3-20 Dec 1985 (stam. 
fl), W. Lewis etal. 10180 (BRIT, MO, USM). 

Cybianthus spr/icei was misinterpreted by Mez, and confused with C. sim- 
plex (Mez 1902). From there, Weigeltia panamensh was described based pri- 
marily on subtle differences and geography. Finally, Cuatrecasas described 
Weigeltia purpurea from the Choco floristic region of Colombia, notable only 
for its narrower leaves, the secondary veins more arcuate, and some quanti- 
tative floral characteristics. 

Cybianthus spnicei is most closely related to C. simplex because of the homomerous 
flowers, long petioles and non-mucronulate leaf apices. Flowever, Cybianthus 
sprucei may easily be separated from C simplex because of its arborescent 
habit, pyramidal panicle with spicate branches, symmetric leaf base and longer 

19. Cybianthus simplex (Hook, f ) G. Agostini (Fig. 4C), Acta Biol. Venez. 
10: 163. 19H(). Comomyrsine simplex Hook, f /« Benth. & Hook., Gen, PI. 2:644. 
1876. Weigeltni simplex (Hook, f ) Mez in Engl., Pflanzenr IV. 236(Hefr 9): 290. 1 902. 
Tvpt: ECUADOR. Chimborazo: At foot of Volcan Chimborazo, 760 m, Aug I860 
(pist, fl, fr), /?, Spri/ee 61 43 (holotyph: K-2 sheets). 

Weigeltia chamaephytci Diels, Nonzbl. Bot. Gart. Berlin-Dahlem 15:383. 1941. syn. nov. 
Cybianthus chamaephyta (Diels) G. Agostini, Acta Biol, Venez. 10:163. 1980. Type: 
ECUADOR. Pasta/.a: Mera, 1,200 m, 15 Nov 1938 (stam. fl), H. Schnltze-Rhonhof 
2983 (HOLOTYPH: B-destr. ; no isotype found). I defer neotypification until material 
has been regathered from the type locality or an adjacent one. 

Senii-woody siibshrub to 1 (—3.5) m tall. Stem terete, 1—5 cm diam., sparsely 
glandular-papillate, glabrescent. Cataphylls few, spirally arranged in inter- 
nodal areas, coriaceous, subulate, 6.5—26.5 mm long, 1.12—1.7 mm wide, 
keeled, densely and prominently black punctate and punctate-Iineate, sparsely 
glandular-papillate, the margin flat, entire. Lfir;z'^.s"pseudoverticillate; blacies 
membranaceous, elliptic to oblong, (34.5— )38— 46. 5(— 80) cm long, (12. 5-) 

PiPOi.Y, Cybianthus in Ecuador and Peru 81 

13. 5- 18. 5 (-3 2. 5) cm wide, apicaliy broadly acute or rounded to a short 
acumen, rarely acute, basally asymmetric, broadly rounded or rarely abruptly 
acute, slightly decurrent on the petiole, hydropotes few above, numerous 
below, often sparsely glandular-papillate below, conspicuously black punc- 
tate, the margin irregular, hyaline when juvenile, opaque at maturity, flat, 
entire; petioles rigid, deeply canaliculate, 7-1 7 (-2 1) cm long, 4-6 mm diam., 
slightly marginate at junctions of the blade, expanded basally and slightly 
decurrent on stem, sparsely glandular-papillate, glabrescent. biflorescence bracts 
membranaceous, linear-lanceolate, 17-25 mm long, 2-4 mm wide, apicaliy 
long-attenuate, hyaline, densely and prominently black punctate-lineate, 
the margin irregular, entire; peduncle ( 1 .2-)2.4— 4.5 cm long. Staminate inflorescence: 
a thyrsoid panicle (columnar) 1 1-28(-54) cm long, bi- or tripinnate, the 
primary branches subopposite, each branch pyramidal-paniculate, the flowers 
racemose; inflorescence branch bracts linear-lanceolate, subulate, (50-)6l- 
72 mm long, 0.5-1.3 mm wide, apicaliy long-attenuate, medially keeled, 
hyaline, glandular-papillose throughout, the margin entire; floral bracts early 
caducous; pedicels 2.3-3(-5) mm long, glandular-papillate and -ciliolate. 
Staminate flowers 5-merous, green to greenish-white when fresh; calyx mem- 
branaceous, subcotyliform, 1.1-1.4 mm long, equally divided, the tube 0.2 
mm long, the lobes lanceolate to lanceolate-subulate, apicaliy narowly acute 
to long-attenuate, hyaline, epunctate, the margin erose-dentate; corolla coriaceous, 
rotate, 2.2-3.1 mm long, the tube 0.3-0.7 mm long, the lobes linear-lan- 
ceolate, 1.5-2.6 mm long, 0.9-1.1 mm wide, reflexed distally 180° from 
the tube, the apicaliy subulate, densely glandular-granulose throughout within 
and along the margin, epunctate, the margin regular, entire; stamens 1.0- 
2.3 mm long, the tube conspicuous, coriaceous, 0.3—0.7 mm long, the apicaliy 
free portions of the filaments terete, as wide as the anthers, 1.0-1.1 mm 
long, ventrally recurved, the anthers connivent at first, subglobose, 0.4- 
0.5 mm long, 0.5-0.6 mm wide, apicaliy rounded, basally cordulate, dorsifixed 
just above the base, the connective prominently black punctate dorsally; 
pistillode absent. Pistillate inflorescence: as in staminate but 16-30 cm long; 
inflorescence branch bracts ovate-subulate, 4.5-7 mm long, 1.5-2 mm wide, 
the margin irregular, erose; floral bracts membranaceous, linear-subulate, 
1.6-2 mm long, 0.4-0.6 mm wide, apicaliy long-attenuate, densely glan- 
dular-papillose puberulent, the margin irregular-entire; pedicel terete, 1- 
10 mm long, translucent, glandular-puberulent. Pistillate flowers green; ca- 
lyx coriaceous, 0.6-1 mm long, unequally divided, the tube 0.2-0.3 mm 
long, the lobes ovate to widely ovate, 0.4-0.7 mm long and wide, apicaliy 
acute, often moderately glandular-papillate without, the margin irregular, 
erose; corolla 1.5-1.7 mm long, the tube 0.4-0.5 mm long, the lobes ob- 
long, 1.0-1.2 mm long, 0.7-0.8 mm wide, apicaliy acute, reflexed 135° 

82 SiDA 18(1) 

from tube, glandular-papillate without; staminodes 0.8-1.2 mm long, the 
tube inconspicuous, membranaceous, 0.4-0.5 mm long, glabrous, elobate, 
the apically free portions of the filaments 0.3-0.4 mm long, the antherodes 
subglobose, 0.2 mm long, 0.3 mm wide, apically rounded, basally cordulate; 
pistil subglobose, 0.8-1.1 mm long, the ovary 0.6-0.8 mm long, 1.0-1.2 
mm diam., densely translucent-lepidote, the style short, thick, 0. 1 mm long, 
0.7 mm diam., the stigma capitate, the margin lacinate, with numerous 
lobules to 0.2 mm long, the placenta umbonate, the ovules 3, exposed apically 
1/3 their length on the placenta. Fruit subglobose, 4-5 mm long, 5-8 mm 
diam., red, exocarp somewhat thick, juicy, prominently black punctate. 

Distribution. — Endemic to the slopes of the Western Cordillera of the Andes 
in Colombia and Ecuador, 60-2,200 m. 

Ecology and conservation status. — Cybianthus simplex occurs in premontane 
wet and rainforests, occasionally in the understory of ridgetops forests. This 
species occurs in deep shade under the shrub stratum of the forest. It occurs 
more frequently in primary forest wet enough to maintain Chusquea popu- 
lations, but can survive in disturbed forests as long as the shrub and Chusquea 
populations exist. Because of its habitat flexibility, it is not considered threatened 
or endangered. 

Etymology. — The epithet "simplex" refers to the monoaxial habit of the 

Specimens examined, COLOMBIA. Nariiio: Mpio. Barbacoas, C^orregimiento Orn'z y 
Zamora, Vereda El Barro, Reserva Natural Rio Nambi', ca. 5 km W de Aitaquer, faldas 
occidentales de la Cordillera Occidental, 01° 18' N, 78° 08' W, 1,350-1,400 m, 1 Sep 
1997 (scam, fl), /. Pipoly, A. Cogollo. d al. 2!02{^ . 21046, 2105L 2]055_{BR1T, FMB, 
JAUM, PSO), 2 Sep 1997 (bisex. (Dj. P,po/y. A. Co^ollo et al. 2/7(;9_(BRIT, JAUM, PSO), 
(ster.)J. Pipoly. A. Coi^ollodal. 21 131. 21148 (BRIT, PSO), (pist. fl, fr),/ Pipoly. A. Co^ollo 
etal. 21173 (BRIT, FMB, JAUM, PSO), 1, -190-1, 500 m, 4 Sep 1997 (stam. fl),J. PipoIy, 
A. Cogolloetal. 21294 (BRIT, FMB, JAUM, PSO), (pist. fl),/. PipoIy. A. Cogolloetal. 21296 
(BRIT, FMB, JAUM, PSO), 1,350-1 ,400 m, 5 Sep 1997 (scam, fl), /. Pipo/y. A. Cogo/Io et 
al. 21326 (BRIT, JAUM, PSO), (ster.),J, Pz/Wj. A. Cogollo ,t al. 21328 (BRIT, JAUM, 
PSO),l, 350-1, 145 m, 6 Sep 1997 (ster.),J. Pipoly. A. Cogolloetal. 21417 (BRIT. FMB, 
JAUM). 1 Sep 1997 (fr),/ Pipoly. A. Cogolloetal. 21469. 21471, 1,450-1,500 m, 8 Sep 
1997 (scam, fl),/ Pipoly. A. Cogolloetal. 27,520 (BRIT, FMB, JAUM, PSO), (ster.),/. Pipoly. 
A. Cogollo et al. 21524 (BRIT, JAUM, PSO); along trail from main Pasco-Tumaco Rd. to 
Rk) Nambi, departing main Rd. ac Escuela Mixta El Mirador, 7 km W of Alcaquer, 01" 
18' N, 78° 04' W, 1,100 m, 26 Feb 1992 (fr), T. Croat 72394 (JAUM, MO); Corregimiento 
Alcaquer, Vereda el Barro, Reserva Nacural Rfo Nambf, W slope, W Cordillera, 01° 18' 
N, 78° 08' W, near Cabana Fundacion FELCA, 1,325 m, 1 1 Dec 1993 (fr),/. Betancur et al. 
4857 (COL, MO); La Planada, Finca Salazar, 7 km above Ricaurce, 01° 08' N, 77° 58' W, 
1,750 m, 29 Nov 198 1 (pist. fl, fr), A. Ge>/try etal. 35188 (BRIT, COL, MO, US); La Planada, 
SofRicaurte, 7 km from Tumaco-Pa.sto Rd., 01° 10' N, 77° 58' W, 1,800 m, 24 Jul 1986 
(scam, fl), A. Gentry et al. 55053 (MO, PSO); trail to Hondon, 6-12 km SW of La Planada, 
01° 04' N, 78° 02' W, 1,750-1,800 m, 5 Jan 1988 (fl bud), 0. de Benavu/es & R. Keating 
6041 1 (MO, PSO); Valley of Rfo Guiza, Rd. from El Espino to Tumaco, ca. 21 km W of 

PlPOLY, Cybianthus in Ecuador and Peru 83 

Ricaurtc, 01° 1 5' N, 78" 07' W, 1 ,000 m, 7 Dec 1 988 (stam. fl), B. Hamniel 17150 (JAUM, 
MO). ECUADOR. Bolivar: Hacienda Changuil, LA 16; 02° 06' S, 79° 10' W, 500 m, 17 
Aug 1995 (stam. H), X. Cornejo & C. Bonifaz 4339 (GUAY, MO). Carchi: Prominent hillcrest 
directly N of Lita, on N side of Rio Mira, E of Rfo Baboso, W-facing slope, 00° 5.^' N, 78° 
27' W, 760 m, 7 Aug 1994 (ster.), B. Boyle 347 3 (MO, QCNE), Steep N-facing slope S of 
Baboso, S side of Rio Baboso, 00° 53' N, 78° 27' W, 750 m, 1 1 Aug 1994 (ster), B. Boyle 
3399 (MO, QCNE); Ri'o Blanco drainage above Chical, tributary of Rfo San Juan, 12 km 
W of Maldonado, 1300-1500 m, 25 Sep 1979 (bisex. fl), A. Get/try & C. Shupp 26363 
(MO, QCNE); Canton Tulcan, Parroquia Tobar Donoso, Reserva Indigena Awa, Centro El 
Baboso, 00° 53' N, 78° 25' W, 1 ,800 m, I 7-27 Aug 1992 (stam. fl), G. Tipaz et al. 1709 
(BRIT, MO, QCNE), (stam. fl), G. Tipciz el al. 1H86 (MO, QCNE), (ir), G. Tipaz et al. 1924 
(MO, QCNE); 6 km above Maldonado, just below Puentc de Palo, 00° 54' N, 78° 06' W, 
2,275 m, 23 May 1993 (stam. fl), B. Boyle &J. Bradford 1878 (MO, QCNE); Trail from 
Pailon to Gualpi Chico, Reserva Indi'gena Awa, 1 .5 km past Rfo Blanco, 00° 5 1 ' N, 78° 
16' W, 1,000-1,450 m, 14 Jan 1988 (stam. fl), W. Hoover et al. 2436 (MO, QCNE); SE 
Trail, Gualpi Chico area of Awa Reserve, 00° 58' N, 78° l6' W, 1,330 m, 19 Jan 1988 
(pist. fl, fr), W. Hoover et al. 2809 (MO, QCNE); Trail along ridge and forest slope to NW 
of Awa encampment, Gualpi Chico area near Finca Rodriguez, 00° 58' N, 78° 16' W, 1,258- 
1 ,323 m, 19 Jan 1988 (fr), W Hoover et al. 3338 (MO, QCNE). Cotopaxi: Rio Guarapa, ca. 
20 km NW of El Corazon, 250 m, 19 Jun 1967 (stam. fl), B. Sparre 17091 (S), 20 Jun 
1967 (pist. fl), B. Sparre 17081 (S). El Oro: 1 1 km W of Las Pinas on new Rd. to Sta. Rosa, 
850 m, 8 Oct 1979 (stam. fl), C. Dodsoii et al. 9101 (MO, SEE); Hacienda Buenaventura, 
12 km W of Las Pinas on Rd. to Machaia, 03° 48' S, 79° 46' W, 1,000 m, 1 Mar 1991 
(stam. fl), M. Kessler2601 (GOET, MO); New Rd. Saracay-Balzas-Velacruz, ca. 8 km SE of 
Saracay, 400 m, 30 Apr 1980 (stam. fl), G. Harling & L. Andersum 18778 (GB). Guayas: 
Cordillera Chogon-Colonche, Cerro Los Pontones; 01° 44' S, 08° 40' W, 500 m, 2 Jul 1994 
(stam. fl), X. Cornejo & C. Bonifaz 2979 (GUAY, MO). Loja: Tierra Colorada, I km E of 
Landara, 8kmEofMercadillo,04°02'S, 79° 57' W, 1,500 m, 9 Feb 1991 (pist. fl, fr), M. 
Kessler 2401 (BRIT, GOET). Manabi: Machalilla National Park, zona de San Sebastian, 
01 ° 36' S, 80° 42' W, 600-700 m, 2 1 Jan I 99 1 (fr), A. Gentry et al. 72499 (MO, QCNE). 
Pichincha: Quito-San Juan Chiriboga-Sto. Domingo de los Colorados Rd., Branch km 
59, 18 km NW of Rd., 1,700-2,000 m, 27 Sep 1 986 (bud), V. Zak 1330 (MO, US); Quiro- 
Aloag-Sto. Dominigo de los Colorados, km 94, 10 km S of Rd., W slopes ol Volcan El 
Corazon, 00° 21' 30 "S, 78° 51' 15" W, 1,300-1,500 m, 25 Dec 1986 (fr), V. Zak 134') 
(MO, US); 15 ha. Patch of forest in Cooperativa Sta. Marta No. 2, along Rio Verde, 2 km 
SE of Sto. Domingo de Los Colorados, 530 m, 5 Feb 1979 (fr), C. Dodson et al. 7397 (MO, 
SEL); Reserva Flon'stica-Ecologica "Rfo Guajalito," Km 59, Quito-Santo Domingo de los 
Colorados, 3.5 km NE of Rd., lower slopes of Volcan Pichincha, 00° 13' 53" S, 78° 48' 10" 
W, 1 ,800-2,200 m, 28 Dec 1985 {k)J.Jaraniillo 8298 (MO, QCA); Canton Quito, Parroquia 
Nanegal, Reserva Maquipucuna, along Inca Trail to Rfo Tulambf, ca. 5 airline km SE of 
Nanegal, 00° 07' N, 78° 38' W, 1 ,350 m. 1 5 Sep I 989 (fr), G. Webster & P. Delprele 27394 
(DAV, QCA), along trail between Rfo Umachaca and Rfo Tulambf, 00° 07.5' N, 78° 38.5' 
W, 1,200-1,300 m, 7 Jul 1990 (fr), G. Webster et ai 27793 (DAV, QCA); Montarias de 
Maqmpucuna, Cerro Sosa, 00° 0.5' N, 78° 37' W, 1,950 m, 3 Jul 1991 (fr), G. Webster 
28702 (DAV, QCA), 1,750 m, 3 Jul 1991 (stam. fl), G. Webster et al. 28710 (DAV, QCA), 
on ridge between Base Camps 1 & 2, 00° 5.5' N, 78° 37' W, 1,800-1,900 m, 6-7 J til 
1991 (fr), G. Webster & B. Castro 28769 (DAV, MO, QCA); along Rfo Umachaca near Ha- 
cienda El Carmen, 00° 07-7.5' N, 78° 38' W, 1,250 m, 6-7 Jul 1991 (fr), G. Webster etal. 
28796 (DAV, QCA). Quininde: Bilsa Biological Reserve, Montarias de Mache, 35 km W 
of Quininde, 5 km W of Sta. Isabela, SE ridge trail, 00° 21' N, 79° 44' W, 400-600 m, 21 

84 SiDAlcSd) 

Sep 1994 (scam, tl), N. Pitiiuiii c/ al. 688 (MO, QCNE), 5 Dec 1994 (fr), N. P/lm/i/ 993 
(MO, QCNE), Alon^ Dogala and Invaders Trails, 00° 21' N, 79° 4i' W, 4()()-6()() m, 2 
Jan 1995 (fr), N. P/liinin I I6l (MO, QCNE). Province unknown: withoLir locality, Sep 
1896 (pist. fl),./, Soc/iro 1U()/12 (B-descr., F Neg 4859). 

This species has often been confused with CyhianthHS ipri/cei, owing to 
variation in leaf morphology and inflorescence size. However, recent field 
studies conducted at the Rfo Nambi Natural Reserve of Narifio, Colombia, 
have revealed that juvenile individuals have obtuse to somewhat broadly 
rounded, asymmetric leaf bases with long petioles, while mature individu- 
als have tapering, asymmetric leaf bases. The confusion was due, in part, to 
precociously flowering individuals, detectable by their extremely small flowers, 
or to reiterative shoots, detectable by the renewal shoot visible below the 
"bayonet", that bears juvenile leaves and pink flowers. While the largest 
individuals of Cybianthus siniphx may approach the size of many C sprncei. 
the large pith of the stem in the former renders them extremely weak, and 
the stems may easily be snapped by hand, while the pith of C. spriwei is 
relatively smaller, and the stems can be bent without snapping in the field. 

Cybianthiis sniipkx is most closely related to C. sprncei, but may be sepa- 
rated from it by the columnar, thyrsoid panicles with racemose branches, 
the pedicellate flowers, asymmetric leaf base and shorter cataphylls. Popu- 
lations corresponding to the type of Weigeltia cbcinuiephyta differ from the 
type of C. simplex in floral structure, directly attributable to the fact that 
the former is based on a pistillate, and the latter a staminate collection. However, 
the autapomorphic columnar thyrsoid panicle leaves no doubt that they are 
synonymous. Because no further collections have been made in the region 
from which the type was collected, I defer neotypification until collections 
from that area are available. 

20. Cybianthus kayapii (Lundell) Pipoly, comb. nov. (Figs. 3E, 4F, 6J, 

9A-F). \Vi7c;(-///)/ kjyapi! Lundell, Writ;htia 6: 118. 1 980. Tvfh: PERU. Amazoxas: 
Camino de chichijam, cntsa, 300 m, 2 May 1973 (fr), R. Knyap 723 (i iolotyfi:: MO; 
isoTYPi;: LL-TEX). 

Monoax'u/l treelet to l(-2) m tall. Stems terete, (O.(S-)l-l .7 cm diam., glandular- 
papillate at first, glabrescent. Cataphylls alternate in a high spiral, coria- 
ceous, subulate, 1 5—45 mm long, ().5-2(-3.5) mm wide. lu^ives psetidoverticillate; 
blades chartaceous, oblanceolate to oblanceolate-oblong, (22-)31.3-55.5 
cm long, 8.5-19.6(-23) cm wide, apically acute or broadly rounded to a 
small acutish ti]^, mucronate, the mucron to 0.5 mm long, the blade gradually 
tapering to an abruptly obtuse base appearing auriculate, to 1.5 cm wide, 
midrib slightly raised above, prominently raised below, the secondary veins 
12-16 pairs, with prominent marginal and submarginal collecting veins, 
slightly sunken above, prommently raised below, glabrous above, with ru- 
fous hydropoten below; petiole deeply canaliculate, 1-2 cm long, ca. 3-5- 

PiPOi.Y, Cybianrhus in Ecuador and Peru 85 

4 mm diam., densely glandular-papillate adaxially. Staminate inflorescence: a 
pyramidal, bipinnate panicle, (3-)5.5-29 cm long, 5-15(-26) cm wide, the 
branches racemose, densely glandular-papillate, succulent, then drying hyaline; 
peduncle 3-5.5 cm long; branch bracts membranaceous, subulate, 6.5-8 
mm long, 0.5-1.5 mm wide; pedicels cylindrical, 1.2-1.8 mm long, sparsely 
glandular-papillate, glabrescent; floral bracts membranaceous, subulate, inserted 
on the pedicel about at middle, longer than the pedicel, 1.5-2.5 mm long, 
0.1-0.2 mm wide, hyaline, densely glandular-papillate, the margin entire. 
Staminate flouws pink, heteromerous, the calyx 5-merous, the corolla 4-merous; 
calyx deeply membranaceous, cupuliform, 0.9-1. 1 mm long, the tube 0.1- 
0.2 mm long, unequally divided, the lobes deltate to subdeltate, 0.6-0.9 
mm long, 0.2-0.7 mm wide, highly reflexed at anthesis, apically acute, epunctate, 
hyaline, densely glandular-papillate, the margin glabrous, entire; corolla 
carnose, subrotate to rotate, 2-3 mm long, the tube 0.5-0.8 mm long, the 
lobes ovate, 1.5-2 mm long, 1.1-1.6 mm wide, apically acute, distally re- 
curved 90° from tube axis at anthesis, opaque, densely glandular-granulose 
within and along margin, sparsely glandular-papillate along margins without, 
epunctate or sparingly and inconspicuously pellucid punctate, the margin 
entire; stamens 2.2-2.9 mm long, subequal to corolla lobe or exserted, the 
tube conspicuous, carnose, 0.5-0.8 mm long, hyaline, glabrous, elobate, 
the apically free portions of the filaments terete, 1 .6-2.2 mm long, free from 
corolla, proximally recurved, the anther oblate, 0.3-0.5 mm long, 0.5-0.8 
mm wide, always wider than long, apically emarginate to retuse, basally 
widely cordate, the connective prominently black punctate dorsally, con- 
spicuously black punctate ventrally; pistillode absent or to 1 mm long, 0.1- 
0.3 mm wide, densely glandular-papillate. Pistillate inflorescence as in stami- 
nate but 6.5-9(-10.5) cm long, erect, not succulent, opaque, densely 
glandular-papillate; peduncle 1-2 cm long; branch bracts 2-3 mm long, 
0.2-0.3 mm wide; pedicels subobsolete or cylindrical, to 1 .2 mm long, incrassate 
and accrescent in fruit to 1.5 mm long; floral bracts inserted on pedicel, 
longer than the pedicel, 1-1.3 mm long, 0.2-0.3 mm wide. Pistillate flowers 
as in staminate but pink to pinkish-white; calyx 0.9-1 .2 mm long, the lobes 
unequally divided, the smaller linear-lanceolate, 0.8—0.9 mm long, 0.3— 
0.4 mm wide, the larger deltate, 1.0-1.1 mm long and wide; corolla ro- 
tate, 2.6-2.9 mm long, the tube 0.9-1 mm long, the lobes elliptic, 1.7- 
2.0 mm long, 0.6-0.7 mm wide, reflexed at anthesis, distally rectirved 180° 
from tube axis, glabrous without, sparsely glandular-granulose within, the 
margin slightly irregtilar; staminodes very poorly developed, 1.2—1.5 mm 
long, the tube 0.9-1.0 mm long, the apically free portions of the filaments 
0.3-0.6 mm long when developed, recurved proximally, the anthers badly 
formed, at times consisting of 2-3 prominently punctate glands at filament 
apex, or orherwise as in the stamens, ovate to subglobose, 0.2-0.3 mm long, 

86 SiDA 18(1) 

0.3-0.4 mm wide, apically irregular, obtuse, emarginate or refuse, basaliy 
cordate, the connective when distinguishable prominently black punctate 
dorsally, conspicuously punctate ventrally; pistil clavate to lageniform, 3- 
3.5 mm long, the ovary 0.9-1 . 1 mm long, 1 .2-1 .5 mm diam., densely papillate, 
the style 2. 1-2.4 mm long, the stigma large, capitate, with 4 principal lobes, 
each irregularly lacinate-lobulate, early caducous, the placenta deeply cupuliform, 
the ovules 2, buried for 1/2 their length. Frai/ depressed-globose, 5-6 mm 
long, 7-9 mm wide, prominently black punctate, the exocarp thin. Bisex/uil 
inflorescence: as in stammate but 4-13 cm long. Bi sex// a I flowers as in stami- 
nate flowers but calyx 1.1-1.9 mm long, the tube 0.2-0.3 mm long, the 
lobes unequally divided, deltate to elliptic, the smaller 0.6-0.7 mm long 
and wide, the larger 0.9-1.6 mm long, 0.5-0.6 mm wide, otherwise as in 
pistillate flowers; corolla 2.6-2.8 mm long, the tube ca. 0.6 mm long, the 
lobes narrowly ovate, 2.0-2.2 mm long, 1.2-1.3 mm wide, recurved dis- 
tally 90° Irom tube, sparsely glandular-granulose within, glandular-papil- 
late along the margin; stamens as in staminate flower, but 2.2-2.7 mm long, 
always slightly shorter than corolla tube, the tube ca. 0.6 mm long, the 
apically free portions of the filaments 2.0-2.2 mm long, the anthers widely 
ovate, ca. 0.3 mm long, 0.5-0.6 mm wide, apically obtuse to emarginate, 
basaliy widely cordate; pistil 2.7-2.9 mm long, the (wary 0.8-0.9 mm long, 
0.6-0.7 mm diam. , densely glandular-papillate, the placenta deeply ctipuliform, 
the ovules 2, buried for 1/2 length. B/id^A7/f7//r///y depressed-globose, 5-6 
mm long, 6-7 mm wide, prominently black punctate, the exocarp thin. 

D/str/b//tion. — Colombia (Amazonas, Choco, Narino, Valle del Cauca, ), 
Ecuador (Chimborazo, Napo and Pichincha), Peru (Amazonas, Loreto, San 
Martm) and Brazil (Acre), growing at sea level-2,530 m elevation. 

Ecology cind couservatio}/ stat/ts. — Cyb/anth/is kayap'// is locally common in 
small populations at the high watet line in primary "tahuampa" habitats 
(varzea forest), along white water rivers, or rarely in premontane habitats 
along the edge of pools led by creeks. The species is not known to be culti- 
vated and occurs only in areas where deep leaf litter and alluvial deposits 
are left after flooding action. It appears that the species can easily be de- 
stroyed by soil compaction as a result of trampling, and thus, it should be 
considered threatened. 

Etymology. — The epithet commemorates Rubio Kayap, an indigenous 
Aguaruna Peruvian plant collector who worked with Brent Berlin, known 
for his great knowledge of Amazonian flora and ethnobotany. 

Local names and /ises. — Colombia: "Hierba de palo grande"(Spanish). Ground 
in crude syrups to cure cuts, internal infections, to clean the uterus and the 
stomach. Ecuador: "putush" (Shuar). Used against intestinal parasites and 
for chronic rectal bleeding (colo-rectal carcinoma °); 10 lbs. of root boiled 
in 8 liters of water, down to one liter; 8 cc given as enema before bed; useful 

PiPOLY, Cybianthus in Ecuador and Peru 87 

for "dysentery." Peru: "mantaya," "kugkuima muspari" (Aguaruna); used 
to disinfect dog and insect bites; "napi tsuake" (Huambisa). Brazil: used in 
curare cf. B. Krukoff7663. 

Representative specimens examined. COLOMBIA. Amazonas: Mpio. Leticia, Parque 
Nacional Natural Amacayacu, Centre Administrativo Mata-mata, trail to Amacayacu, km 
4, 03° 47' S, 70° 15' W, 120 m, 25 Sep 1991 (stam. fl), A. Ruclas & A. Pneto5l47 (COL, 
FMB, MO), 110-120 m, 28 Oct 1991 (A bud),/ Ptpoly &J. Murillo 15483 (COL, FMB, 
MO), Quebrada de Agua Pudre, ca. 1.5 km NE of junction with Ri'o Amacayacu, perma- 
nent inventory plot, 200-220 m, 11 Nov 1991 (ster.),/. Pipoly et al. ] 5896 (COL, FMB, 
MO), 15 Nov 1991 (stam. fl),/ Ptpoly etal. 16075 (COL, FMB, MO); Rio Loretoyacu, 100 
m, Oct 1946 (bisex. fl), R. E. Schultes & G. Black 8427 (US). Choco: Bahia de Solano, IS- 
IS Apr 1982 (pist. fl, fr), R. Dressier 6036 (COL, FLAS, MO); Mpio. de Quibdo, Corregimiento 
San Francisco Icho, Quebrada Caledonia along Caledonia Rd., 9 Apr 1987 (ster.), F. Garcia 
&J. Echavavria 259-A (COL, CHOCO, MO). Narino: Mpio. Ricaurte, Reserva Natural La 
Planada, 1,800 m, 13 Nov 1993 (fr), C. Restrepo 723 (BRIT, MO, PSO); La Planada, Finca 
Salaazar, 7 km above Ricaurte, 01° 08' N, 77° 58' W, 1,750 m, 27 Nov 1981 (fr), A. Gen- 
try et al. 35062 (MO, PSO). Narino: Mpio. Barbacoas, Corregimiento Ortiz y Zamora, 
Vereda El Barro, Reserva Natural Rio Nambi, ca. 5 km W de Altaquer, faldas occidentales 
de la Cordillera Occidental, 01° 18' N, 78° 08' W, 1,350-1,400 m, 3 Sep 1997 (ster.),./. 
Pipoly, A. Cogollo, et al. 21241 (BRIT, FMB, JAUM, PSO). Valle del Cauca: Rfo Calima, 
region del Choco, La Trojita, 5-50 m, 20 Feb 1944 (stam. fl),/. Cuatrecasas 16272 (COL, 
F, US); Mpio. El Cairo, Cerro del Ingles, summit, Cordillera Occidental, Serrania de los 
Paraguas, limit Vallc/Choco, El Cairo-Rio Blanco Hwy, 1 hour in jeep from El Cairo, 2,400 
m, 1 Jan 1987 (pist. fl), P. Silventom-Sopktn et al 2871 (CUVC). ECUADOR. Carchi: Canton 
Tulcan, Parroquia Tobar Donoso, Reserva Indigena Awa, Centro El Baboso, 00° 53' N, 
78° 25' W, 1,800 m, 17-27 Aug 1992(stam. R),G. Tipazetal / 706 (MO, QCNE). Esmeraldas: 
Canton San Lorenzo, Parroquia Ricaurte, Reserva Indfgena Awa, Comunidad Balsarefio, 
Rfo Palabf, 01° 09' N, 78° 31' W, 100 m, 15-29 Apr 1991 (fl bud), D. Ruhio & C. Quelal 
1335 (MO, QCNE). Chimborazo: Cordillera Occidental, "El Carmen," Sibambe, 2,450 
m, 22 Aug 1943 (pist. fl), Al, Acosta-Solh 5544 (F, QCNA); on slopes of Chimborazo Vol- 
cano, (pist. fl, fr), A. Sodiro 100114 (B, destr., QA?, n.v.). Morona-Santiago: Basin of Ri'o 
Morona, Rfo Mangosiza, Nayumbime, 45 km SE of Sucua (by air), 200 m S of Don Lui's 
Najamte's house, ca. 02° 43' S, 77° 38' W, 300 m, 27 Feb 1990 (fr), C. Limbach 140 (QCA, 
NY). Napo: Antisana, Shinguipino Forest, between Rios Napo and Tena, 8 km SE of Tena, 
01° 00' S, 77° 50' W, 450 m, 17 Sep I960 (stam. fl), P. Griibhetal. / 633 (K, NY); Canton 
Archidona, Carretera HoUin-Loreto, Rio Huataraco, 2 hrs by foot from Guagua Sumaco, 
00° 43' S, 77° 32' W, 800-1 ,000 m, 23-30 Aug 1989 (fl bud), C. Ceron & M. Eactos 7641 
(MO, QCNE); Canton Tena, 17 Oct 1939 (stam. fl), E. Asplund9396 (S); 3 km E of Caseri'o 
Huamanf, N of Carretera Hollin-Loreto, 00° 43' S, 77° 36' W, 1,200 m, 17 Sep 1988 (fr), 
F. Hi/rtado & A. Alvarado 503 (MO, QCNE). Pichincha: Along Rd. from Tandayapa to 
Mindo, 10 km from Tandayapa, 2,530 m, 16 Dec 1979 (fr), T. Croat 49361 (MO, QCNA). 
PERLJ. Amazonas: Prov. Bagua, Dtto. Imaza, Comunidad Aguaruna de Kampaentza (PUJ AIM), 
property of Juan Mayan, 740 m, 6 Oct 1994 (pist, fl), C. Diaz et al. 7265 (BRIT, HUT, 
MO, USM); Prov, Condorcanqui, Dtto. El Cenepa, NE region of Marahon Drainage Basin, 
Rfo Cenepa, Comunidad Tutino, 04° 33' S, 78° 10' W, 350 m, 21 Nov 1993 (pist. fl, fr), 
R. Vasq/tez et al. 18395 (AMAZ, BRIT, HUT, MO, USM); Rfo Cenepa, vicinity of Huampami, 
ca, 5 km E of Chavez Valdfvia, 04° 30' S, 78° 30' W, Quebrada Chigkishinuk, 10 Apr 
1973 (fr), E. Anaiash 211 (MO), 11 Aug 1978 (pist, fl), E. Ancuash 1405 (MO); Trail one 
day walk from Huampami to Shaim, creek running into Nahim, 600 m, 27 Nov 1972 


SlHA 18(1) 

Fic;. 17. Cybianthjis anthnriophyll//s Vipoly. A. Habir, showint; pleiochasium wich sympo- 
dial branches. B. Sraminate flower, sliowing lanceolate-SLibulatc corolla lobes, elobare scaminal 
tube, and conic, vestigial pistillode. C. Pisrillace flower, showing attenuate-acicular calyx 
lobes and obturbinate pistil. A, C, drawn from holofype. B, drawn from Bravo & Gomez 49. 
Figure drawn by Peggy Duke. 

PiPOLY, Cybianthus in Ecuador and Peru 89 

(scam. H), B. Berlin 395 (MO); Rio Santia^-o, W bank, 400 m beyond La Poza, 180 m, 
without date (fr), F. Donii'ng/iez 147 (MO); 800 m beyond Caterpiza, 200 m, 4 Sep 1979 
(fr), V. Hiiasbikat 3^6 (MO), 10 Sep 1979 (stam. fl), V. Hiiaihikat 507 (MO), 1 2 Sep 1979 
(bisex. ID, V. Huashikat 5H1 (MO); 26 Mar 1 980 (fr), S. Tunqiii 1110 (MO). Loreto: Prov. 
Maynas, Explorama Lodge, near Yanamono, between. Indiana and mouth of Rfo Napo 03° 
28' S, 72° 50' W, 103 m, 27 Jun I 983 (fr), Gei//)y el al. 42247 (AMAZ, MO, NY), 106 m, 
15 Apr 1985 (fr), R. Vasquez & N.Janiwillo 6325 (AMAZ, MO), 28 Sep 1988 (stam. fl), R. 
Wdsqmz & N.Jaramillo 11100 (AMAZ, MO)l 1 Jul 1990 (fr), R. Vdsqmz & N. Jciraiiiillo 
14092 (AMAZ, MO, US), 25 km NE of Iquitos, along Rfo Amazonas, 90 m, 03" 30' S, 
72" 50' W, 26 Sep 1990 (stam. fl),./, Pi/m/y vt al. 12383 (AMAZ, MO), along S border 
trail, 1 10 m, 27 Sep 1990 (stam. i\),J. P/poly ct al. 12490 (AMAZ, MO, NY, US, USM), 
(scam. fl),J. Plpoly et al. 12541 (AMAZ, MO, US, USM); Prov. Alto Amazonas, N slopes of 
Cerros Camapaqufz at Pongo de Mansenthe, right bank of Ri'o Maraiion, 300-550 m, I 9- 
21 Oct 1962 (stam. fl),J. Wurdack 2324 (US, USM). San Martin: Pongo de Cainarachi, 
Rio Cainarachi, tributary of Rio Huallaga, 230 m, Sep-Oct 1962 (stam. fl-except speci- 
men at S-bisex. fl), G. King 2691 (A, F, GH, K, MO, NY, S, US). BRAZIL. Acre: Cruzeiro 
do Sul, Rios Jurua and Moa, 8 km above Cachoeira Grande, 07° 30' S, 73° 30' W, 27 Apr 
1971 (fr), G. Prance etal. PI 2555 (IAN, MG, NY). Amazonas: Basin of Riojavarf, Mpio. 
Sao Paulo de Oliven<,-a, near Esperanga, Dec 1935 (fr), B. Kri/koff7663 (NY, U). 

Cybianthm kayapii may be confused with C. sprucei (Hook, f.) G. Agostini, 
but may be recognized by the shorter petioles, heteromerous flowers and 
mucronate leaf apices. However, the leaf blades with mucronulate apices 
and subauriculate bases, and heteromerous flowers indicate Cybianthus kayapii 
is more closely related to C. anthiiriophyllm. From Cybianthus anthrmophyllus, 
C. kayapii may be separated by the much smaller, flat, entire leaves with 
entire margins, the deltate calyx lobes with entire margins, and the carnose 

21. Cybianthus anthuriophyllus Pipoly, sp. nov. (Fig. 17). Type: ECUA- 
DOR. Napo: Canton Gonzalo Pizarro, Rfo Tigre, affluent of Rfo Dashino, entering 
from 73 km of Rd. from Lumbaque to El Reventador, 10 km S oi Lumbaque, 00° 
05' S, 77° 24' W, 900-1,100 m, 18-21 Feb 1 987 (pist. fl, fr), W. Palacios & D. Neill 
1584 (hoi.otype: US; i.sotypes: K, MO, QCNE). 

Ob folia oblanceolata ab lamina ad pctiolum gradatmi contracco, a primo intuitu cum 
C kayapi confusa est, sed ab ea marginibus laminaribtis revolutis serrulatisque (nee integettimis 
planisque) inflorescentiis 28 (non 8-15) cm longis, pleiochasia cum ramulis floriteris cymosis 
(nee bipinnatipaniculatis cum ramulis floriferis racemosis) lobulis calycinis subulatis (nee 
ovatis) praeclare distat. 

Monoaxial treelet to 0.6 m tall. Stems terete, weakly woody, ca. 2 cm diam., 
sparsely glandular-papillate, glabrescent. Cataphylls tightly pseudoverticillate, 
coriaceous, linear-subulate, 4.0-6.5 cm long, 0.3-0.6 cm wide, densely and 
prominently punctate-lineate, sparingly glandular-papillate, glabrescent. 
Leaves pseudoverticillate, erect; blades coriaceous, narrowly lanceolate, (56-) 
104-1 10cm long, (l4.5-)17-22 cm wide, apically broadly rounded or rounded 
to a minute abrupt submucronate acumen 2 mm long, the blade gradually 
tapering to the petiole, almost obtusish basally, midrib slightly raised above. 

90 SiDA 18(1) 

prominently raised below, the secondary veins ca. 42 pairs, impressed above, 
prominently raised below, nitid and epunctate above, purple (when fresh), 
inconspicuously punctate and papillate-puberulent below, the papillae erect, 
rufous, the margin thin, opaque, inroUed, prominently straight-serrulate, 
the teeth alternatively larger and smaller (ca. 1 mm and 0.5 mm long, re- 
spectively); petiole deeply canaliculate, thick, ca. 1.5 cm long, 0.8 cm diam., 
sparingly glandular-papillate. Staminate inflorescence: a pyramidal thyrsoid 
panicle, ca. 16 cm long, 23 cm wide, the branches pseduoracemose (sym- 
podial), the rachis densely glandular-papillate, the peduncles longer below, 
shorter above; inflorescence branch bracts linear-lanceolate, 7—9 mm long, 
2-3 mm wide, apical ly attenuate-acicular, sparsely glandular-papillate, the 
margin entire; floral bracts acerose, 2-3 mm long, 0.4-0.8 mm wide; pedicels 
1.0-2.5 mm long, angular, thin, the longer pedicels ui the lower portion of 
the inflorescence, the shorter ones above, densely glandular-papillate, the 
papillae persistent. Staminate flowers heteromerous, membranaceous, hyaline, 
the calyx (5-)6-merous, the corolla 5-merous; calyx subcotyliform 3.7-4.7 
mm long, 0.4—0.7 mm wide, apically acuminate-acicular, prominently keeled, 
the keel thickened hyaline, epunctate, sparsely glandular-papillate except 
densely so along the margin, the margin entire; corolla subrotate, 7.4—10 
mm long, the tube 0.7-0.9 mm long, the lobes lanceolate-subulate, 6.3-9 
mm long, 2.7-3.2 mm wide, the apically long-acuminate, highly reflexed, 
sparsely glandular-papillate without, very sparsely glandular-granulose within 
above the junction within tube the margins densely glandular-papillate, 
entire; stamens 2.2—4.1 mm long, the staminal tube conspicuous, coria- 
ceous, 1.4-1 .8 mm long, glabrous, elobate, the apically free portions of the 
filaments ventrally recurved, 0.8-0.9 mm long, flat, glabrous, the anthers 
ovate, 0.3-0.4 mm long and wide, apically and basally emarginate, dorsifixed 
less than _ length, the connectives prominently black punctate ventrally 
and dorsally; pistillode conic, vestigial-0.8 mm long or absent. Pistillate 
inflorescence a pleiochasium, with branches cymose (sympodial), the rachis 
densely glandular-papillate, the peduncles 1-3 cm long, longer below, shorter 
above; inflorescence bracts resembling cataphylls but smaller, 1.3-1.6 cm 
long, 1 .0-1.3 mm wide, conspicuously punctate-lineate; inflorescence branch 
bracts linear-lanceolate, 1 .2-1 .5 mm long, 0.2-0.4 mm wide, apically at- 
tenuate-acicular, sparsely glandular-papillate, the margin entire; floral bracts 
acerose, longer than the pedicels, 0.5-1.0 cm long; pedicels angular, thin, 
1.2—7.5 mm long, the longer pedicels in the lower portion of the inflorescence, 
densely glandular-papillate, the papillae persistent. Pistillate flowers as in 
staminate but calyx 6-merous, (2.1-)3-4.1 mm long, the tube 0.3-0.7 mm 
long, 0.5-1 .0 mm wide, apically long-attenuate-acicular; corolla (from dried 
remnants) subrotate, hyaline, 5.0-7. 1 mm long, the tube 1 .0-1 .3 mm long, 
the lobes hnear-subulate, 4.0-5.8 mm long, 1.1-1 .5 mm wide at base, apically 

PiPOi.Y, Cybianthus in Ecuador and Peru 91 

subulate, highly reflexed, twisted and distally recurved at anthesis; stami- 
nodia 1.2—1.3 mm long, the staminodial tube 0.9—1.2 mm long bearing 
well-developed lobes alternate with the apically free filaments, the filaments 
0.3—1.1 mm long, flat, glabrous, the antherodes malfi^rmed, mostly con- 
sisting of undifferentiated tissue surrounding prominent black punctations 
or suborbicular and 0. 1—0.5 mm long, 0. 1—0.6 mm wide, always wider than 
long, apically emarginate, dorsifixed slightly less than _ length, the connectives 
prominently black punctate dorsally; pistil obturbinate, 2.5—3.0 mm long, 
the ovary 1 .0—1.5 mm long, 0.8-1.3 mm wide, densely papillate and prominently 
black punctate, the style 1.3-1.7 mm long, densely glandular-papillate, the 
stigma large, capitate, lobes, the lobes lacinate, each lobe to 0.4 mm long, 
early caducous, the placenta widely conic, bearing 4 uniseriate, exposed ovules, 
the ovules on the periphery of the placenta. Bisexual fruit pink, globose, 4— 
6 mm long and diam., the punctation prominent, brown when fresh (teste 
coll.), red or black upon drying, the exocarp thin. 

Distribution. — Amazonian ("Oriente") Ecuador and adjacent Peru (Loreto), 
l60-300(-l,100) m elevation. 

Ecology and conservation status. — Cybianthus anthuriophyllus grows in pri- 
mary tropical wet forest and premontane wet forest on terra firnie above the 
high water contour. It is found in primary forest as well as in secondary, but 
it is not known whether the plant is cultivated in secondary forest situa- 
tions. At this time, the species can be considered locally common but not 

Etymology. — The specific epithet refers to the unique shape of the adult 
leaves, held erect in vivo and reminiscent o{ Anthurium crassinerviuyn (Araceae). 

Local names and uses: — Ecuador: "namakuk" (Achuar JiVaro); "Acuari" (dialect 
unknown); "challuo panga" (Quichua). Peru: "kutukupish," "takushia," 
"mutupash," "kurup" (Maynajivaro), "sierra panga" (Quichua). Leaves crushed 
and used as a fish poison( W. Leivis et al. 14051); inner stem is scraped and 
an infusion given to dogs to drink to improve their hunting abilities and to 
enhance their stamina {W. Lewis et al. 1 1 133): stem is scraped and its juice 
put into a dog's nose to produce sneezing {W. Lewis et al. }()475), or to "make 
it an attack dog against thieves" (Lewis et al. 12833). 

Paratypes. ECUADOR. Napo: Cuyabeno-Punta AreniUa; Sep 1981 (stam. fl), E. Bravo 
&P. Gamez 49 (QCA); Canton Orellana, Yasunf National Park, Maxus Rd. and pipiineconstmction 
project km 10, 00° 29' S, 76° 34' W, 250 m, 29Jun 1994 ( fr), N. Pitwati 448 (JVtO, QCNE). 
Pastaza: Kapawi (Amuntaf), rfo Pastaza; Village area, 02° 31' S, 76° 48' W, 235 m, 25- 
29 Jul 1989 (ster.), W. Lewis et at. 14051 (MO). PERU. Loreto: Prov. Alto Amazonas, 
Puranchim, Rio Sinchiyacu, 02° 50' S, 76° 55' W, 200 m, 3-7 Dec 19<S8 (stcr.), W. Lewis 
et al. 14390 (MO); Washintsa and vicinity, Rfo Huasaqa, 03° 20' S, 76° 20' W, 185 m, 
16-26 Jun 1986 (ster,), W. Lewis et at. 11153 (MO); Prov. Loreto: Nueva Jerusalem and 
vicinity, Rio Macusari, 02° 55' S, 76° 15' W, 220-300 m, 29 Dec 1985-3 Jan 1986 (Fr), 
W. Lewis et at. 10475 (MO); Pampa Hermosa and vicinity, Rio Corrientes, 1 km S of junc- 

92 SiDA 18(1) 

cion with Rio Mucusari, O.V 15' S, 75" 50' W, 16() m, 3-20 Dec 19H5 (sram. H), W. Leins 
et cil. lO.UO (MO); Vista Aiegre, Rio 'I'igre, 02° 40' S, 75° 35' W, 210 m, 17 Mar 1987 
(stcr.), W. Leir/s ci a/. 1285,^ (MO). 

Cy/jh///tb//s ai/thur/ophyII//s is unique within subgenus Cnmoni^niue by its 
small, marginal pectinate leaf serrations and pleiochasial inflorescence, ap- 
pearing paniculate, but with sympoclial primary and secondary branches, 
and thus, cymose. The leaves appear subsessile, with the blade gradually 
tapering to the deeply canaliculate petiole, a feature found otherwise only 
in its closest congener, Cyhianthus kayapii. However, Cybianthns anthuriphyllin 
is clearly distinct from C. kaycipii because of the inroUed, pectinate-serru- 
late leaf blade margin, tlie much longer, pleiochasial inflorescence, and subulate 
calyx lobes. The extremely long cataphylls are the best developed in the 

V. Cyhianthus subgenus Triadophora (iVIez) G. Agostini, Acta Biol. Venez. 
1 0: 1 64. 1 9H(). Wasi^dtn-t subgenus TridJophoni Mez in Engl., Pflanzenr. IV. 236(Hei'c 
9):291 . I 902. Typi; Species. Wcii^dtid sdtliiiiii (Hook, f.) Mez in Engl. = Cyh'hnithi/s 
schl/niii (Hook, i.) G. Agosrini. 

Currdiunid D'Arcy, Ann. Missouri Hot. Card 60:442. 1973. Type Species. Corrdlnvui 
spectcihilis (Standi.) D'Arcy = Cyhu/iithus .uhliinii (Hooker 1.) Ct. Agostini. 

As here interpreted, Cyhictnthns subgenus Tnculophora is monotypic. Its 
only species, C. schlni/ii, is easily recognized by its monoaxial habit and 
autapomorphic rufous glandular tomentum of malpigiaceous trichomes, leaf 
blades with subepidermal fibers and pseudocataphylls (here defined as petiolate 
cataphylls). The first full description oi Cyhianthi/s schliiini is provided, along 
with complete synonymy and complete exsiccatae for Ecuador and Peru, 
and representative ones for other areas. 

22. Cyhianthus schlimii (Hook, f.) G. Agostini (Fig. 3D, 6K, 6L), Acta 
Biol. Venez. 10:165. 1980. Connwiyrsiiw schliwii Hook. f. in Benth. 6- Hook., 
Gen. Pi. 2:644. 1 876. Weigdtui sddunii (Hook. \.) Mez in Engl., Pflanzenr. IV. 236(Heft 
9):29l. 1902. CoynUuina Khlimn {W^oV. f.) D'Arcy, Ann. Missouri Bot. Cjard 60:443. 
1973. Type: COLOMBIA. Meta: Llano de San Martin, 300 m, Jan 1836 (stam. fl), 
J. 'Wunui 7594 (iioi.oi ype: K; esotypes: COL, LE, MA, P). 

Weigeltici multijlora A.C. Smith., Bull. Torrey Bot. Club 60:387. 1 933- syn. nov. CmreUiiinu uiiiU't- 

flora {K. C. Sm.) D'Arcy, Ann. Missouri Bot. Gard 60:4 15. 1973. (A. C. Sm.)G. Agostini, 

Acta Biol. Venez. 1 0: 165. 1980. Type: BRAZIL. Mato Grosso: Near Tabajara, upper 

Rio Machado, 2t Nov 1931 (sram. fl), B. A. Kri/koff 1 .]HS (hoi.otyi'i,: NY; lsotype: A). 

Aniisni ipeclahilis SvAnd[., Publ. Eield Mus. Nat. Llist. Bot. Ser. 18:893. 1938. syn. nov. 
Weigdtia spectabilLs (Standi.) Lundell, Wright ia 4: 1 69. 1 97 1 . Cotrdliami speclahilis (Standi.) 
D'Arcy, Ann. Missouri Bot. Gard. 60:443. 1973- Cyhunith//.s spaUihilis (Standi.) G. 
Agostini, Acta Biol. Venez. 10:165. 1980. Type;: COSTA RICA. Aeajuiea: Cataratas 
(Los Angeles) de San Ramon, A|-ir 1935 (stani. ti), /\, Bn//a 20"^ iO (molotype: E; esotypes: 
CR, F). 

Wc'/gdt/j trunidni Aspl., Bot. Not, I9t9:8{)2. 1939. T\m- COLOMBIA. Cauca: Near 
Distrito El Tarn bo, 900 m, 3 1 Jul 1936 (stam. II), K. vtm SiiL'/der)i 919 (i ioeo'EYPi:: S). 

PiPOi.Y, Cybianthus in Ecuador and Peru 93 

Weigeltid schiimii (Hook, f.) Mtv van inteniiecl/n Moklcnkc, Phytokigia 2:242. 1 947. Typi;: 
COLOMBIA. Vai.ii; di;i. Calica: Pacific coa.s:, Rfo Cajambre, San Isidro, 5-100 m, 2- 
5 May 1944 (stam. fl),_/. Ci/citreaiscis 1 7312 (noioTVi^r,: NY; isotypi-s: COL, F-2 sheets). 

Monoaxtal tree to 5 m. Stem terete, 0.8—2.0 cm diam., the wood dense, 
minutely rufous glandular appressed tomentose, the trichomes malpighiaceous, 
early glabrescent. Pseiidocataphylls produced only irregularly, chartaceous, 
subulate, ca. 2.0—3.0 cm long, 0.5—1.0 cm wide, apically acute, mucronate, 
densely rufous puberulent, black lineate-punctate, the margin entire; peti- 
ole subobsolete, to 0.2 cm long. Leaves tightly pseudoverticillate; blades 
chartaceous, elliptic, oblong or oblanceolate, 25—65 cm long, 5.5-20 cm 
wide, apically acute or subacuminate, mucronulate, the mucro often sclerified, 
the acumen 0.5-3.0 cm long, base long-attenuate, the blade decurrent on 
the upper portion of the petiole, midrib slightly elevated above, prominent 
below, the secondary veins 9—13 pairs, prominent, the marginal veins loop 
connected, conspicuously striolate by subepidermal fibers, these visible above 
and below, sparsely rufous puberulent above, moderately puberulent be- 
low, at times glabrescent, hydropotes absent, sparsely to densely punctate 
or lineate-punctate below, the margin opaque, irregular, entire to roughly 
serrate; petioles canaliculate, 1 .0— 3.0(-10) cm long, 0.5—1 .0 cm diam., abruptly 
swollen basally, puberulent, glabrescent. Staniinate Inflorescence a pinnate or 
bipinnate columnar panicle 13^0 cm long, 3-20 cm wide, the rachis densely 
glandular-papillate and rufous puberulent, the flowering branches racemose; 
peduncle 8-15 cm long; inflorescence bract chartaceous, ovate, 9—15 mm 
long, 2.4—4.5 mm wide, apically acute, densely rufous glandular puberu- 
lent, conspicuously black punctate and lineate-punctate, the margin opaque, 
entire; inflorescence branch bracts membranaceous, linear, 10—13 mm long, 
1.9—2.1 mm wide, apically narrowly acute, mucronulate, minutely rufous 
puberulent, orange furfuraceous lepidote, densely and conspicuously black 
lineate-punctate, the margin opaque, entire; floral bracts membranaceous, 
subulate, 0.8—1.3 mm long, 0.2—0.4 mm wide, sparsely rufous puberulent, 
the margin entire; pedicel terete, 2.0^.5 mm long, prominently black punctate, 
densely papillate and rufous puberulent. Staminate flowers 3(^)-merous, light 
purple, then dull yellow; calyx chartaceous, cupuliform, 0.9-1-8 mm long, 
the tube 0.3 mm long, the lobes subdeltate, 0.7—1.6 mm long, 0.9-1-2 
mm wide, apically acute, densely rufous puberulent, glabrescent, densely 
and prominently black punctate, the margin flat, wide, hyaline, densely ciliolate, 
the cilia often caducous; corolla rotate, chartaceous, 3.0-5.0 mm long, the 
tube hyaline, 0.8—1.0 mm long, the lobes elliptic or oblong, 2.2—4.0 mm 
long, 1.6—2.2 mm wide, apically obtuse to rounded, subcucullate, invo- 
lute, distally recurved 180" relative to tube, sparsely rufous puberulent without, 
very sparsely glandular-granulose within basally, often glabrescent, very densely 
and prominently black punctate except margin hyaline, irregular, glabrous, 

94 SiDA 18(1) 

entire; stamens 2.4-3.5 mm long, the tube membranaceous, inconspicu- 
ous, 0.8—1.0 mm long, hyaline, epunctate, glabrous, elobate, the apically 
free portions of the filaments 1 .8—2.3 mm long, sparsely or epunctate, sparsely 
rufous puberulent at first, glabresccnt, the connective prominently punc- 
tate, or inconspicuously so, the punctation orange, red or black, the anthers 
cordate, 0.5-0.6 mm long, 0.7-1.0 mm wide, apically subacute to rounded, 
base deeply cordate, dorsifixed at point less than 1/5 distance from apex; 
pistillode none or conic, to 1.0 mm long, densely and prominently black 
punctate and rufous papillate. Pistillate inflorescence as in staminate but more 
columnar, (3.5-)8.0-] 8.5 cm long, (2.0-)3.5-6.() cm wide, the branches 
subcorymbose to rarely racemose; peduncle 1.5—6.8 cm long; inflorescence 
bract ovate to elliptic, 4.5-12 mm long, 1.9—4.0 mm wide, conspicuously 
black punctate; inflorescence branch bracts 5-15 mm long, 0.5-2.5 mm 
wide; floral bracts 0.9-2.0 mm long, 0.3-0.6 mm wide; pedicels terete, 
2.0— 4.0 mm long, accrescent to 5.0 mm long and incrassate to 2.0 mm 
diam. in fruit. Pistillate flowers as in staminate but 3-merous, calyx purple, 
corolla yellow; calyx subcotyliform, 1.8-2.0 mm long, the tube 0.2-0.3 
mm long, the lobes subdeltate to widely ovate, 1.6-1.8 mm long, 1.6-2.2 
mm wide, apically acute, the margin slightly erose, conspicuously long glandular- 
ciliate, the cilia often caducous; corolla 4.2—5.0 mm long, the tube 0.3-0.5 
mm long, the lobes oblong, 3-2-4.5 mm long, 2.3-2.6 mm wide, apically 
obtuse, somewhat cucuUate, reflexed ca. 45" from tube axis, densely pu- 
berulent without, sparingly glandular-granulose basally within; staminodes 
to 2.5 mm long, the tube membranaceous, inconspicuous, to 0.5 mm long, 
the filaments adnate to corolla lobe ca. 0.5 mm long, then apically free to 
1.0 mm long, thick, terete, rarely punctate, the antherodes cordate, ca. 0.6 
mm long, 0.7—0.8 mm wide, apically acute to apiculate, base widely cor- 
date, the connective punctate, prominently or not; pistil clavate to obnapiform, 
4.0—5.2 mm long, the ovary 1.8 mm long, 0.5-1.6 mm wide, prominently 
black punctate and rufous puberulent, the style to 1.3 mm long, densely 
rufous puberulent, the stigma capitate to lobed, 3(— 4)-lobed, the placenta 
deeply cupuliform, bearing 2 ovules exposed apically. Print globose, 1.0— 
1.3 cm long, 1.0—2.0 cm diam., at maturity, the exocarp bright orange to 
red-orange, prominently black punctate. 

Distribution. — From the Atlantic Slope (Dpto. Ri'o San Juan) Nicaragua, 
to state of Pando, Bolivia and adjacent Amazonia of Brazil, 0-1,800 m. 

Ecology and conservation status. — Cyhianthus schlimii occurs in a variety of 
habitats, from lowland to premontane tropical moist, wet and pluvial for- 
est. It is locally common, but restricted to primary forest. Therefore, it should 
be considered threatened. 

Etymology. — The species is named for Louis Joseph Schlim, a Belgian plant 
collector working for J.J. Linden in Brussels, who collected extensively around 

PiPOLY, Cybianthus in Ecuador and Peru 95 

Caracas to the Venezuelan Andes, and from there, to the grasslands of Meta 
and the Sabana de Santafe de Bogota, Colombia, during the period 1841- 
1852. Schlim also collected with Nicolas Funck later in Venezuela. 
Local names and uses. — Peru: "napi tsuake" (Huambisa). 

Representative specimens examined. NICARAGUA. Rio San Juan: El ReJos, midpoint 
between El Castillo and Delta de San Juan, 0-50 m, 23 Mar 1961 (stam. fl), G. Bunting & 
L. Lick 77^ (F, NY). COSTA RICA. Alajuela: 15 km NW of Arenal by air, 2 km NW of 
Nuevo Arenal on Rd. to Tilaran, then 3 km NE on Rd. to San Rafael de Guatuso, then 2 
km W on Rd. to Finca Cote, 10° 34' N, 84" 54' W, 700 m, SE side of Lago Cote, 30 Apr 
1983 (fr), R. Liesmretal. 15093 (CR, MO, WIS). Heredia: Zona Protectora, N slopes of 
Volcan Barba. betweem Rio Peje and Rio Guacimo, along Quebrada Cantarana, 300-400 
m, 18 Jan 1983 (stam. fl), M. Grayum&G. 5/)^/z 3 i 70 (CR, DUKE, MO); Finca La Selva, 
Rio Puerto Viejo 2 km E of jet with Rio Sarapaqui, 10° 26' N, 84" 00' W, 100 m, 14-17 
Jun 1968 (stam. fl), W. Burger & R. Stolze 5803 (CR, F, MO, NY). Limon: Near Finca 
Castilla, 30 m, 24 July 1936 (ster), C. Dodge & V. Goerger9283 (F, MO). San Jose: Estacion 
Carrillo, Cafion del Rio Sucio, 450-700 m, 12 Nov 1983 (pist. fl), /. Chacon & G. Herrera 
1720 (CAS, CR, MO, NY). PANAMA. Bocas del Toro: Cerro Bonyic, above Quebrada 
Huron, 180-400 m, 13 Apr 1968 (fr),/ Kirkhnde &J. Duke 610 (MO). Colon: Base of 
Cerro Bruja, along Ri'o Escandaloso, above Mina Boqueron, No. 2, 47.5 km from Tran- 
sisthmianHwyonRd. to Salamanca, 09° 50' N, 79° 32 'W, 10-200 m, 18 Mar 1982 (stam. 
fl), S. Knapp & W.J. Kress 4282 (MO, NY, PMA). Darien: Rio Tuquesa, Tuquesa Mming 
Co. camp, Charco Peje, 250 m, 7 Jul 1975 (stam. fl), 5. Mori 7015 (MO, SCZ). Panama: 
Cerro Campana, 800 m, 22 Jun 1967 (fr), T. Croat 17167 (MO). San Bias: Trail from Rio 
Estadi to Cerro Banega, 300-530 m, 09" 23' N, 78" 51' W, 21 Dec 1985 (stam. fl), G. de 
Nevers & H. Herrera 6642 (CAS, MO, PMA). COLOMBIA. Antioquia: 6 km E of Guapa, 
53 km S of Turbo, 240 m, 13 May 1945 (stam. fl), 0. Haught 4660 (US); Mpio. Anori, 
Corregimientode Providencia, Buenos Aires, 500-600 m, 4 Feb 1972 (fr), D. Soejarto 3205 
(HUA, MO, NY); Vicinity Planta Providencia, 26 km S, 23 km W (by air) of Zaragoza, 
07" 13' N, 75" 03' W, valley of Rio Anori between Dos Bocas and Anorf, 1 Jun 1976 (fr), 
J. Shepard323 (COL, WIS); Vicinity Medellin, 20 Aug 1927 (stam. fl), R. Toro 356 (MEDEL, 
NY); Medellfn-Bogota Hwy, sector Rio Samana-Ri'o Claro-San Luis, 400-1,000 m, 24 Aug 

1982 (fr),/ Hernandez & S. Hoyos 483 (COL, HUA); Mpio. Urrao, Boundary of Parque 
Nacional Natural Las Orqufdeas, Vereda Calles, Permanent Inventory, Premontane Rainforest, 
left bank of Rio Calles, 06° 32' N, 76° 19' W, 1,450-1,500 m, 30 Nov 1993 (ster.),/ 
Pipo/y et al. 17406 (BRIT, JAUM, MO). Boyaca: Region of Cerro Chapon, extreme W 
part of Boyaca, NW of Bogota, 2,300 m, 31 Jul 1932 (fr), A. Laurence 370 (A, NY, S); El 
Umbo region, 1,000 m, 12 Oct 1932 (stam. fl), A. Laurence 530 (A, BM, F, G, GH, MO, 
NY, S, U, UC, US). Choco: Mpio. Quibdo, Quebrada La Platina, Hwy to Medelli'n, 25 Sep 

1983 (fr), L. Arias eta/. 134 (MO); Mpio. San Jose del Palmar, along Rfo Torito (affluent of 
Rio Habita), W slopes, 850- 950 m, 15 Mar 1980 (fr), E. Forero et al. 7350 , 16 Mar 1980 
(fl bud), E. Forero et al. 7393 (COL, MO), Vereda Portachuelo, Hacienda Barro Blanco, 1 ,350 
m, 15 Jan 1983 (fr), P. Franco et al. 1325 (COL); Rio Mecana, ca. 10 km E of Mecana, 06" 
15'N, 77"25' W, 100 m, 7 Mar 1983 (stam. fl),A. Gentry & A. J iincosa 41072 {QO\.,U.O, 
JAUM), 710-880 m, 8 Jan 1984 (stam. fl), A. Juncosa 1769 (COL, MO, JAUM); Mpio. 
Novita, vereda Curundo, left bank, Rio Ingara, 550 m, 1 Dec 1983 (fr), P. Franco et al. 
1059 (COL); Rio Nuqui, 400 m, 25 Jan 1947 (stam. fl), 0. Haught 5479 (COL, US); La 
Mojarra, upriver from Istmma, 05" 12' N, 76" 37' W, 30-60 m, 5 Nov 1983 (fr), A.J/mcosa 
1255 (COL, JAUM, MO, NY); S of Rio Condoto, between Quebrada Guarapo and Mandinga, 
120-180 m, 22-28 Apr 1939 (fr), E. Ktlltp 35675 (COL, US). Cundinamarca: Cordillera 

96 SiDA 18(1) 

Bo^'oni, 2,000 m, Sep 1H55 (Fr),/. 'I'm/i/a 4 (BM), Meta: Near Salitre, 6 Jan 1876 (stam. 
ti), E. A/Jc/n 1 151, 7 Jan 1876 (ster), E. Andre s.n. (K); C]ano Tigre, between Cano Aguas 
Claras and Cano Grande, 4.5 km SW of ViUavicencio, 04" 07' N, 73" 39' W, 500-550 m, 
24 Feb 1943 (pist. H), ER. Eoiherg 20148 (COL, US); Llano de San Martfn, (scam, fl, pisr. 
fl mixed), f/. Karstc/i s.n. (LE-2 sheets). Nariiio: Mpio. Barbacoas, C^orreginiiento Orci'z y 
Zamora, Vereda El Barro, Rcserva Natural Rfo Nambi, ca. 5 km W de Altaquer, laldas 
occidcntales de la Cordillera Occidental, 01° 18' N, 78" 08' W, 1,350-1,400 m, 1 Sep 
1997 (stam. fl), /. P/po/y. A. Cogollo. et al. 2/095 (BRIT, FMB,JAUM, PSO), 21144, 21149. 
1,450-1,500 m, 2 Sep 1997 (fr),./. P//mIy. A. Cogollo et ul. 21483 1250-1,350 m, 8 Dec 
1997 (ir),y, P/po/y. A. Cogollo et cd. 21598 (BRIT, JAUM, PSO). Norte de Santander: 
Ocana, I 200 m, May 1846-52 (stam. fl), L. Schlim ()86 (BR-3 sheets, F-2 sheets, G-BOIS- 
2 sheets, G-DEL, MA-2 sheets). Santander: 8 km SE ot Barrancabermeja, Ri'o Opon, 200 
m, 28 Atig 1954 (stam. fl), R. Cdstiinedci 4~^4(^ (COL), vicinity Barrancabermeja, between 
Ri'os Sogamoso and Colorado, 100—500 m, S of Rio Sogamoso, Camp Mesa, 8 Jan 1935 
(stam. fl\ 0. Haiight 1502 (A, COL, F, NY, US). Valle del Cauca: Cordillera Occidental, 
W slope, along Rio Sanc]Liininf, left side, La Laguna, 1,250-1,400 m, 13 Dec 1943 (stam. 
fl),J. Citdtyecdias 15474 (VALLE); Bahfa Malaga, Quebrada Algeria, new Rd. to military 
base,0 4" 02' N, 77" 22' W, 50 m, A. Gentry et al. 53326 (COL, MO, US); Bank of Rfo 
Digua, Rfo San Juan, below Queremal, to the right ol river between km 52 and 53, 1300— 
1500 m, 19 Mar 1947 (fr),/ Ctuttrecma.^ 23855 (COL, F-3 sheets); Calima Dam, 1600- 
1700 m, 17 Sep 1966 (stam.).V. Esf>tm/1 2 109 (MO, VALLE). Without locality, date, (stam. 
(\)J. Mittis449 (MA). 29/9 (MA), 3907. 5/02^/ (MA, US), Warscewiczs.n. (B-destr., Photo- 
F Neg. 4858). ECUADOR. Carchi: Trail along plain above Tovar-Donoso and Rio Guape, 
01" 10' N, 78" 18-31' W, 280-450 m, 18 Feb 1984 (stam. fl), W. Hoover 1194 (MO), SE 
Trail, (iualpi Chico Area of Awa Reserve, near encampment, 00" 58' N, 78° 16' W, 1 ,330 
m, 19 Jan 1988 (stam. fl), W. Hoover et al. 2815 (MO, QCNE), Gualpi Medio Community, 
Awa Reserve, 900 m, 2 I rVIay 1992 (bud), C. Quelal et al. 764 (MO, QCNE) El Pailon, 45 
km below Maldonado, along path to Tobar Donoso, 800 m, 1 Dec 1979 (fr), AL Madison & 
E. Besse 7201 (AAU, F, QCNE, SEE); Canton Tulcan, Reserva Indfgena Awa, Parrocjuia 
Tobar Donoso, sector El Baboso, 00° 53' N, 78° 20' W, 1,600 m, 3 Oct 1991 (A, fr), G. 
Tipaz et al. 260 (BRIT, MO, QCNE), (fr), 3 / 1 (BRIT, MO, QCNE), Centro El Baboso, 00° 
53' N, 78° 25' W, 1,800 m, 17-27 Aug 1992 (fr), G. Tipaz et al. 1950 (BRIT, F, MO, 
QCNE). Esmeraldas: Canton San Lorenzo, Reserva Indigena Awa, Canon del Rfo Mira, 
10 km W of Alto Tambo, Comunidad "La Union," 01° 02' N, 78° 26' W, 250 m, 16-26 
Mar 199 1 (fr), D, Riibto et al. 1262 (MO, QCNE). Sucumbios: Canton Lago Agrio, Reserva 
Cuyabeno, Laguna (Trande, Near NEOTROPIC Cabins, 00° 00' S, 76" 1 1' W, 230 m, 15 
Nov 1991 (stam. fl), W Palacios et al. 9269 (BRIT, MO, QCNE). Zamora-Chinchipe: 
Canton Nangaritza, Valle del Rfo Nangaritza, Miazi, 04° 18' S, 78° 40' W, 1,200 m, 10 
Dec 1990 (pist. fl), W. Palacios 67.34 (BRIT, MO, QCNE), Behind military camp, 04° 16' 
S, 78° 42' W, 970 m, 20 Oct 1991 (stam. fl), W. Palacios et al. 8483 (BRIT, MO, QCNE). 
PERU. Amazonas: Camino de chichijam, entsa, 180-250 m, 2 May 1973 (stam. fl), R. 
Kayap 728 (MO); Valle del Rio Santiago, 03° 50' S, 77" 40' W, Quebrada Caterpiza, 2-3 
km behind Caterpiza, 200 m, 4 Jan 1980 (stam. fl), S. Tiinqi/i 549 (MO). Loreto: Prov. 
Maynas, Guarnicion Pijuayal, near Pebas, 130 m, 7 Sep 1988 9stam. fl), C. Diaz et al. 566 
(MO). Ucayali: Prov. Padre Abad, Boqueron del Padre Abad, 400 m, 20 May 1969 (fr),./. 
Schunke 3068 (F-2 sheets, US, USM). BRAZIL. Amazonas: Mpio. Humayta, on plateau 
between Rio Livramento & Rio Ipixuna, 7-18 Nov 1934 (stam. fl), B. Kr//koff7290 (A, 
NY, S, U); Sfu) Paulo de Oliven^;a, near Esperanc;a, Dec 1935 (ster.), B. Kr//koff7(y63 (K, 
NY), Behind Sao Paulo de Olivenga, 16 Atig 1 973 (fr), E. Lleras et al. PI 73 1 ^ (GB, INPA, 
MG, NY); km 500, Manaus-Humayta Rd., 17 Sep 1980 (stam. fl),5. Loirrieetal. 5.2 (INPA, 

PiPOLY, Cybianthus in Ecuador and Peru 97 

MG, NY). Mato Grosso: Near Tabajara, upper Rio Machado, Nov-Dec 193 1 (sram. fl), B. 
Krukoff 1577 (A, NY). Rondonia: SaoLourenqo, cassiterite mine, 20 km Nof S. Louren^'o 
on Rd. to "A Macisa" Mine, 15 Jul 1979 (scam, fl), C. Calderon et al. 2852 (INPA, US). 
BOLIVIA. Pando: W bank of Rio Madeira, 3 km above Abuna, 13 Nov 1968 (stam, fl), 
G. Prance et al. 8388 (LPB, MG, INPA, NY). 

The "pseudocataphylls" referred to in the description are poorly devel- 
oped and do not occur in a regular phyllotactic spiral as is found in subge- 
nus Comomyrsine. Likewise, their morphology is essentially that of a leaf arrested 
at different stages of development, and as such, do not have a distinctive 

Northern populations from Nicaragua and Costa Rica corresponding to 
the type of C. spectabilis (as Ardhia spectabilis) differ in their smaller flowers 
and anther connectives sometimes eglandular, a feature which is more a function 
of ecotype then anything else. Eglandular anthers and entire leaves may be 
found in very lowland wet forest populations in the Darien of Panama, and 
the Choco of Colombia. 

The type of Weigeltia multifiora A.C. Sm. represents populations whose 
inflorescences are less branched (although the duplicates of the type collec- 
tion vary in that regard) and the largest flowers of the species. They, like 
many of the Panamanian populations, have entire leaves and are otherwise 
inseparable from the type of Weigeltia tnandra Asplund. 

The collections of Kayap from Amazonian Peru are referred to this spe- 
cies, despite their longer petioles, 

VI. Cybianthus subgenus Weigeltia (A. DC.) G. Agostini, Acta Biol. Venez. 
10:156. 1980. Weigeltia A. DC, Trans. Linn. Soc. London, Bot. 17:102. 1834. 
Cybianthus sect. Weigeltia (A. DC.) Miq. /;/ Mart., Fl. Bras. 10:299- 1856. Type Spe- 
cies: Salvadora stirnamensis Spreng. Tent 7. 1828. ^Cybianthus siirinaniensis (Spreng.) 
G. Agostini. 

Polyaxial dioecious or rarely, monoecious shrubs or small trees. Roots posi- 
tively geotropic. Trunk distinguishable, growth dynamics following Rauh's 
Architectural Model (in ours), rarely Corner's Model (not in Ecuador or Peru) 
sensi/ Halle et al. (1978). Bark grey to beige, thick, vertically fissured. Branchlets 
sessile furfuraceous lepidote or rarely, rufous tomentose, often glabrescent 
Cataphylls and pseudocataphylls absent. Leaves alternate, rarely subopposite, 
with minute sessile rufous furfuraceous lepidote scales abaxially. Inflorescence 
a pinnate or bipinnate panicle, rarely a simple raceme; peduncle 0.5-2 cm 
long. Flowers 4- rarely 5-merous; calyx valvate, crenate or rarely entire, gla- 
brous or rarely glandular-ciliolate, punctations red or black, prominent or 
not; corolla rotate to subrotate, the lobes mibricate, glandular-granulose 
only at the junction with the corolla tube, the punctations red or black, 
conspicuous, or rarely, prominent; stamens and staminodes united to form 
an inconspicuous or conspicuous tube, the tube without lobes alternating 

98 SiDA 18(1) 

with the apically free filaments, the filaments one to three times longer than 
the anthers, terete, and recurved proximally, the anthers subglobose, or widely 
ovoid, versatile, apically acute to emarginate, basally widely cordate, de- 
hiscent by wide longitudinal slits, the connectives mostly prominently red 
or black punctate; pistil conic, pyriform or obturbinate, the ovary translu- 
cent glandular-lepidote or glabrous, the style capitate-lobate, the lobes entire; 
pistillode minutely conic, or at times, absent. Fruit globose or depressed- 

Cyhiantbiis subgenus Weigdtia contains approximately 46 species in South 
America and the Caribbean. Five species are known from Ecuador and Peru; 
they are restricted to the lowlands and premontane forests on the lower slopes 
of the Western Cordillera in Ecuador and the Eastern Cordillera in Ecuador 
and Peru. 


I . Inflorescence bipinnately paniculate; petioles canaliculate; staminate Mowers 
with stamens shorter than corolla. 

2. Branchlets terete; leaf blades apically long attenuate-subulate, basally long- 

3. Branchlets 2.5-4 mm diam., minutely rufous-lepidote; leaves 
pseudoverticillate, the blades membranaceous, prominently black punctate- 
lineate below, margins subentire to obtusely serrate; petioles 0.5-1.5 
cm long; calyx membranaceous, 1-1.3 mm long, the lobes obtuse, promi 
nently punctate, glandular-ciliate along the margin 23. C. poeppigii 

3. Branchlets 5.5-6 mm diam., minutely ferrugineous tomentellous; 
leaves alternate, the blades chartaceous, minutely and prominently 
pLinctictdose below, margins entire; petioles 2—3-5 cm long; calyx 
chartaceous, 0.7-1 mm long, the lobes acuminate, epunctate, glabrous 

along the margm 24. C. pseudolongifolius 

2. Branchlets angulate or winged; leaf blades apically acute or short-acumi 
nate, basally acute or cimeate. 

4. Branchlets angulate, 8-10 mm diam., conspicuously rubiginous punctate- 
lineate; leaf blades coriaceous, conspicuously rubiginous punctate-lineate 
below; petioles 2.5-3 cm long; staminate calyx membranaceous, 1.4- 
1.6 mm long, the lobes apically acute; staminate corolla membrana- 
ceous, 2.3-2.5 mm long; stamens 1.8-2 mm long 23. C. vascjuezii 

4. Branchlets vvuiged, (2.5-3-)5-6 mm diam., epunctate; leaf blades 
chartaceous, incons|Ticuously pellucid punctate; petioles 2-2.5 cm long; 
staminate calyx chartaceous, 1-1.2 mm long, the lobes apically 
subacuminate; staminate corolla chartaceous, 1.2-1.3 mm long; sta- 
mens l-l.l mm long 26. C. cenepensis 

I . Inflorescence racemose; petioles marginate; staminate flowers with stamens 

exserred or equalling corolla 27. C. nanayensis 

23. Cybianthus poeppigii Mez in Engl., Pflanzenr. IV. 236(Heft 9):218. 

1902. Tvpi:: PF.RU. Lokito: Prov. Maynas. Tocache, without elevation, without 
date, (stam. (i), /:. Poepp/i^ .\.i/. (ikm.otype: W). 

PiPOLY, Cybianthus in Ecuador and Peru 99 

Wei^dtiii alhiflora A. C. Sm., Amer. J. Bot. 27:546. 1940, syn. nov. Cyhiarithi/s cilhiflori/s 
(A.C. Sm.) G. Agostini, Acta Biol. Venez. 1 0:1 57. 1980. Type: COLOMBIA. Choc6: 
Andagoya, 70-110 m, 20-30 Apr 19.39 (stam. fl), E. Ktllip 33372 (hoi.otype: NY; 
isoTYPES: A, BM, US-2 sheets). 

Cybianthus gentry il^undeW, Wright ia 5:195. 1975, syn. nov. Typi;: COLOMBIA. Ch(x;6: 
Cerro Mali', on border with Panama, 1,200-1,400 m, 17 Jan 1975 (stam. fl), A. Gm- 
try & S. Mori 13709 (holotype: LL; isotypi;: MO). 

Shrub or small tree to 4(-7) m tall. Branchlets thin, terete, 2.5-3.5(-4) mm 
diam., minutely rufons-lepidote. Leaves in loose pseudoverticels; blades 
membranaceous, elliptic, oblanceolate, lanceolate or rarely obovate, (6— )6.8— 
18.5(-24) cm long, (2.2-)3.5-6.5(-8.5) cm wide, apically acuminate to 
subacuminate-attenuate, basally cuneate to acute, not decurrent on the petiole, 
midrib depressed above, prominently raised below, the secondary veins 7— 
1 2(-l4) pairs, rufous lepidote above and below early glabrescent above, tardily 
glabrescent below, sparsely black punctate and densely black punctate-lineate 
below, the margin flat, subentire to obtusely serrate; petioles canaliculate, 
0.5—1.5 mm long, minutely rufous-lepidote. Staminate and pistillate inflorescences 
monomorphic, bipinnately paniculate, somewhat pyramidal, sometimes 
malformed and appearing racemose, 8-15 cm long, 8-10 cm wide, the ra- 
chis densely glandular-papillate, the flowers racemose; inflorescence bracts 
unknown; floral bracts membranous, linear-lanceolate, 1— 2.5(— 3-5) mm long, 
0.3— 0.8(— 1.2) mm wide, apically attenuate, densely glandular-papillate, 
epunctate, the margin glandular-ciliate; pedicels cylindrical, (0.7— )1.6— 1.9(— 
2.5) mm long, densely glandular-papillate. Staminate floivers 4-merous, yel- 
low or yellowish-green; calyx membranaceous, cotyliform, 0.8— 1 .3 mm long, 
the tube 0.2-0.3 mm long, glabrous, the lobes widely ovate to suborbicu- 
lar, (().6-)l mm long, (0.5-)0.6-l mm wide, apically obtuse, prominently 
punctate, the margin hyaline, irregular, erose-serrtilate apically, densely glandular- 
ciliate; corolla membranaceous, rotate, (1.7-)1.8-2.2(-2.6) mm long, the 
tube (0.3-)0.4-0.6 mm long, glabrous without, glandular-granulose within, 
the lobes suborbicular, (0.9-)l .2-1 .6(-2) mm long, (0.8-)l .2-1.7(-2) mm 
wide, apically obtuse, prominently punctate, glabrous without, glandular- 
granulose medially at stamen base within, the margin hyaline, irregular, 
erose; stamens (1-)1.4— 1.6 mm long, the tube inconspicuous, hyaline, (0.3-) 
0.4—0.6 mm long, densely glandlar-granulose within, the apically free por- 
tions of the filaments (0.3-)0.4-0.5 mm long, the anthers ovate-triangu- 
lar, 0.3-0.4 mm long, 0.2-0.3 mm wide, apically rounded, basally obtuse, 
the connective epunctate; pistillode come, 0.3-0.4 mm long, densely ru- 
fous glandular-papillate. Pistillate flowers as in staminate, but calyx 1—1.3 
mm long, the tube 0.2-0.3 mm long, the lobes deltate to widely elliptic, 
0.8-1 mm long and wide, the margin opaque; corolla 2-2.3 mm long, the 
tube, 0.6-0.7 mm long, the lobes 1.4-1.6 mm long, 0.6-0.7 mm wide, 

100 SiDA lcS(l) 

glandular-granulose within above filament junction with tube, the margin 
often erose; staminodes resembling stamens but 1-1.3 mm long, the staminodial 
tube ca. 0.6 mm long, the apically free portions of the filaments to 0.3 mm 
long, the antherodes suborbicular, to 0.2 mm long and wide; pistil obnapiform, 
1.2—1 .3 mm long, the ovary 0.8—0.9 rnm long, 0.3—0.5 mm wide, densely 
translucent-lepidote, the style 0.3—0.4 mm long, the stigma bilobed, the 
lobes decurrent, ca. 0.4 mm long. Fruit depressed-globose, 3.5-4.5 mm 
long, 4.5—6 mm diam. when dried, red, then black at maturity, inconspicuously 
punctate, exocarp thin. 

Distribj/iton. — Known from the easternmost Darien of Panama south to 
Amazonian Peru and Brazil, 100-1,800 m. 

Ecology and conservation status. — Cybianthus poeppigii is a broad ranging 
polymorphic ochlospecies, occurring in primary premontane wet and plu- 
vial forests, from the transition zone with lowland forests, to the transition 
zone to cloud forests. Recent fieldwork in the Cordillera Occidental of Co- 
lombia has shown it is a conspicuous element of primary pltivial premontane 
forests, with a density of approximately 20 individuals per hectare, clus- 
tered mostly along the matgin of the windward side of the forest, and along 
streambanks above the high water level. While locally common, its restricted 
primary habitat verifies its threatened status. 

Etymology. — The specific epithet honors Eduard F. Poeppig (1 798-1868), 
professor at Leipzig, explorer and plant collector, who made numerous valuable 
contributions to our knowledge of the Peruvian Amazon Basin. 

Representativf specimens examined. PANAMA. Darien: Scrrani'a del Darien, Panama/ 
Colombia frontier, Cerro Tacaracuna, Cerro Mali, summit, 1,400 m, 17 Jan 1975 (sram. 
fl), A. Gentry &S. Mon 13665 (LL-TEX, MO-2 sheets), W ridge, Cerro Tacaracuna, 1,800- 
1 ,850 m, 3 1 Jan 1 975 (Ir), A. Gentry &S. Mon 1 4023 (COL, LL-TEX, MO-2 sheets); Serranfa 
de Pirre, Cerro Pirre, above Cana Gold Mine between Ri'os Cana and Escucha Ruido, 1 ,()()()- 
1,310 m, 21 Jul 1976 (stam. fl), T. Croat J 7785 (LL-TEX, MO, NY, PMA), SW ridge 
leading to Alturas de Nique, Panama/Colombia border, 1 , 100-1 ,200 m, 30 Dec 1980 (stam. 
fl), R. Hiirt)iki>i 12401. 12461 (MO). COLOMBIA. Antiociuia: Mpio. De Anon', Corregimiento 
Providencia, Rfo Anori Valley, between Dos Bocas and Anorf, •100-900 m, 24-31 May 
1973 Or), D. Soejc/rto et ul. -f()9(} (A, COL, F, LIUA, MO); Mjmo. Sonson, Rfo Verde region, 
Hacienda "La Soledad," 1 ,430- 1 ,800 m, 2 1 Jan 1 947 (stam . fl ), G, G//t/envz 1 1 86 (E, MEDEL, 
MO, LJC); Mpio. San (;arl()s, Corregimiento Alto de Samana, Vereda Miraflores, 820-900 
m, 15 Jun 1989 (stam. fl), R. fomiei^rj et ul. 30^6 (BRIT, IILJA). Mpio. Urrao, Parque 
Nacional Natural Las Orquitleas, Vereda Calles, Permanent Inventory, right bank, Rfo Calles, 
06° 32' N, 76° 19' W, 1 ,450 m, 27 Nov 1993 (ster.),/ P,poly etui HI 83 (BRIT, JAUM, 
MO), 1,450-1,500 m, 29 Nov 1993 (ster. seedling),/ Pipoly et al. / 7360 (BRIT, JAUM, 
MO). Choco: Mpio. San Jose del Palmar, vereda "El Tabor," 1, 540 m, 18 Jan 1983 (pist. 
fl), P. Prcinco et al. 1469 (COL). Vaupes: Rfo Pacoa (tributary of Rfo Apaporis), 00° 20' N, 
71° 20' W, 300 m, 7-12 Feb 1952 (pisr. fl), R. Schi/ltes & I. Cabrera 1^423 (COL, GH, 
US). ECUADOR. Morona-Santlago: Macuma, 50 km NofMacas, 21 Mar 1973 (pist. fl), 
H. Ugo 3633 (GB); Rfo Cuycs and Boboiza-Gualaquiza Rd., 03° 25' S, 78° 35' W, 800 m, 
1 Nov I 986 (stam. fl), U^ Pataaos ! 466 (MO, US). Napo: Between Tena and Napo. 1 Jan 

PiPOLY, Cybianchus in Ecuador and Peru 101 

1940 (stam. fl), U. Aspli/rid 10209 (S), 7 Jan 19i0 (stam. fl), E. Aspl/nid 10302 (S); Tena, 
400 m, 3 Apr 1935 (stam. fl), Y. Alexia 7206 (NY, UC, US); ReservaBiologicaJatun Sacha, 
8 km from Puerto Misahuallo, right bank, Rio Napo, 01° 04' S, 77° 36' W, 450 m, C. 
Cerdn2585 (MO,QCNE). Pastaza: Mera, 1 ,100 m, (stam. fl), E. Asplund 18717(5), 1,500 
m 29 Dec 1958 (stam. fl), G. Hurling et al. 9764 (GB); Loracachi, on path to Lagartococha, 
01° 38' S, 75° 58' W, 25 May 1980 (fr) J. Jaramillo et al. 30984 (AAU, QCA), 31 May 
1980 (tr), 3 I -5 79 (AAU, QCA). Tungurahua: Ri'o Negro, 1 ,200 m, 1 3 Mar 1980 (pist. fl), 
G. Marling & L. Andersson 17255 (GB); 10 km E of Paquisha, 1,400-1,500 m, 13 Apr 
1985 (stam. fl), G. Marling & L. Andersson 24097 (GB). PERU. Amazonas: Rio Santiago, 
3 km behind Caterpiza, 14 Nov 1 979 (fr), V. Muashikat 1248 (MO, NY); Tingo Maria, 25 
Jul 1940 (stam. fl), E. Asplund 12497 (S). Pasco: Prov. Oaxapampa, Cabeza de Mono, Palcazu 
Valley, 10° 20' S, 75° 18' W, 320 m, 1 1 Jun 1983 (fr), A. Gentry et al. 41880 (MO); Ozuz 
to Rio Lobo, 10° 19' S, 75° 16' W, 400-500 m, 10 May 19H5 (fr), R. Foster & B. d'Achilk 
10013 (F). San Martin: Quebrada de Haquisha (right margin Rio Hualiaga), Prov. Mariscal 
Caceres, Dtto. Tochache Nuevo, 400-500 m, 1 Jul 1974 (stam. fl),^. Schunke 7106 (MO, 
NY). BRAZIL. Amazonas: Near Jurua, Dec 1900 (stam. fl), E. Ule 5160 (HBG). 

Cylnanthus poeppigit was mistakenly placed by Agostini (1980) in subge- 
nus Cybianthus, but the versatile, dorsifixed anthers clearly place it in sub- 
genus Weigeltia. I had reported earlier (Pipoly 1983a) that Cyhtanthus albiflorus , 
its synonym, was closely related to C. lawrencei Moldenke. However, the 
bipinnate inflorescences with racemose flowers, stamens shorter than the 
corolla and usually long petioles indicate that Cyhianthm poeppigii is more 
closey related to C. longtfolius Miq., a vicariant species of the southwestern 
Amazon Basin of Brazil and adjacent Bolivia. The glandular grantiles of 
the staminal and staminodial tubes represent a unique (autapomorphic) character 
state within the genus. 

The holotype oi Cylnanthus poeppigii is staminate, as are those of its taxo- 
nomic synonyms. The type of Weigeltia albiflora {Cybianthus albiflonts) rep- 
resents populations with entire, irregularly margined leaves, but is other- 
wise qualitatively identical with that of C. poeppigi. Likewise, the type of 
Cybianthus gentryi Lundell represents isolated montane populations of the 
Darien/Choco regions, and exhibits more notable lineate-punctations, much 
smaller leaves, and abbreviated inflorescences. According to annotations by 
Killip at US, description of another taxon was at one time contemplated, 
based on the fact that the populations of this species from near Tena, Ecua- 
dor, have roughly serrate leaf margins. 

24. Cybianthus pseudolongifolius Pipoly, sp. nov. (Fig. 18). Type: PERU. 
Pasco: Prov. Oxapampa, Palcazu Valley, Cabeza de Mono, 5-6 km W of Iscosacfn 
10° 12' S, 75° 14' W, 325 m, 13-19 Apr 1983 (fr), D. Smith 3808 (holotype; MO; 
isoTYPEs: US, USM). 

Quoad folia magna chartacea, inflorescentias bipinnatipaniculatas, j^ediceilos cylindricos, 
lobos calycinis ovatos, ca. 1/3 connatos, petiolos canaliculatos C. longijolio arete accedens, sed 
abearamuiis teretibus (non angulatis), 5.5-6 (non 3.5-4) mm diammetris, laminis anguste 
oblanceolatis (non ellipticis vel lanceolatis) desuper sordidis (non nitidis) ad apices longi- 


SlDA 18(1) 

Fic. 18. Lybicnithns p\i:iid(ili)iigijtil}iis Pipoly. A. H;ibi:, showing bipinnate panicles. B. Pe- 
duncle and axillant leaf, showmg canaliculate petiole. C;. Detail of prominently puncticulosc 
abaxial leal surtace. D. Pedicel, calyx and fruit, showing hyaline and erose calyx lobes. A & 
C, drawn from isotype; B ik. D, drawn From holotype. Figure drawn by Linda Ellis. 

PrpoLY, Cybianthus in Ecuador and Peru 103 

attenuacis ct subulatis (nee acutis vel atuminatis) ad bases longi-attenuatis (nee aeutis), 
inflorescentiis 8-13 (non 15-20)cni longis, pedicellis 2.8-3 (nee 0.6-1) mm longis necnon 
fructibus depresso-globosis (nee globosis) acque minute cosatis (nee laevibuss) perfacile diseenda. 

Treelet to 3 m tall. Branchlets terete, 5.5-6 mm diam., densely and mi- 
nutely ferrugineous tomentellous. Leaves alternate; blades chartaceous, nar- 
rowly oblanceolate, (22-)26.5-33(-36.5) cm long, (5-)6.5-8.5 cm wide, 
apically long attenuate, subulate, basally long-attenuate, decurrent on the 
petiole, sordid and glabrous above, pallid, minutely and prominently pel- 
lucid puncticulose and minutely ferrugineous puberulent below, midrib slightly 
raised above, prominently raised below, the secondary veins 13-18 pairs, 
the margin entire, glabrous, flat; petioles canaliculate, (2-)2.5-3(-3.5) cm 
long, glabrous above, minutely ferrugineous puberulent below. Staminate 
inflorescence: unknown. Pistillate inflorescence: a lax bipinnate panicle, 8—13 
cm long, 1.5-4 cm wide, densely feraigineous puberulent, glabrescent; secondary 
inflorescence bracts unknown; floral bracts unknown; pedicels cylindrical, 
2.8—3 mm long, densely ferrugineous puberulent; Pistillate flowers unknown; 
fruiting calyx chartaceous, cotyliform, 0.7-1 mm long, the tube 0.3-0.4 
mm long, the lobes ovate, 0.5-0.7 mm long, 0.5-0.6 mm wide, apically 
acuminate, densely and prominently red punctate, the margin hyaline, erose, 
epunctate, glabrous. Fr///7 depressed-globose, 3—4 mm long, 5-6 mm diam., 
inconspicuously pellucid punctate, minutely longitudinally costate. 

Distribution. — Known only from the type. 

Ecology and conservation status. — Cybianthus pseudolongifolius appears to be 
restricted to the lowland primary forest of the eastern Andean slopes. The 
Oxapampa Province of Pasco is home to numerous Peruvian endemics, and 
it would not be surprising if the species was of extremely limited distribu- 
tion or endemic. With only one specimen known, no determination can be 
made of its true conservation status. 

Etymology. — The specific epithet refers to its general likeness to Cybianthus 
longifolii/s Miq., a lowland black water river species from Amazonian Brazil 
and Venezuela. 

Cybianthus pseudolongifolius is most closely related to C. longifolius, but 
differs by its terete branchlets, 5.5-6 mm in diameter, narrowly oblanceolate 
leaf blades that are sordid above, long-attenuate and subulate apically and 
long-attenuate basally, longer inflorescences, much longer pedicels and depressed- 
globose, minutely costate fruits. The Oxapampa region of Pasco contains a 
number of endemic species and disjunct taxa and as such, is one of the most 
important underexplored areas in Peru. 

25. Cybianthus vasquezii Pipoly, sp. nov. (Fig. 19). Type: PERU. Loreto: 
Prov. Alto Amazonas, Capahuari Norte, 02° 45' S, 76° 25' W, 220 m, 7 Jun 1981 
(stam. fl), R. Vdsciiiez &N.Jc!nimillo 1993 (hoi.otype: MO; isotypes: AMAZ, BRIT, 
F, NY, US, USM). 


SiDA 18(1) 

Fic. 19. Cyb'uinthus vasquezii Pipoly. A. Habit, showing angulate, punctate-lineate branciilet. 
B. Inflorescence branch, showing racemose-glomerulatc floral arrangement. C. Open co- 
rolla. D. Abaxial calyx lobe surface. E. Abaxial leaFsurface, showing minute furufuraceous 
scales and prominent punctate-lineations. R Branchlet apex. A-F, drawn by Linda HUis, 
from holotype. 

PiPOLY, Cybianrhus in Ecuador and Peru 105 

Propter ramulos crtissos manifesce angulatos, fol ia coriacea subter pallida ad apices subaciiminata 
ad bases acuta, petioles canaliculatos, inflorescenrias anguste bipinnatipaniculatas, ramulos 
inflorescentiares dense spicatos vel glomerulatos lobos calycines grosse crenatos C. potiaeo 
valde affinis sed ab ea ramulis aibiginoso-puncrato-lineatis (non epunctatis), folia pseudoverticillata 
(non alterna), laminis oblanceolatis (nee ellipticis), (9— )10— 13(— 1 5.3) (nee 6.5—8) cm latis, 
petiolis 2.5—3 (nee 1.5—2) cm longi.s, inflorescentiis 12—25 (non 3—6) cm longis, corollae 
lobis ovatis (non ellipticis) ad apices acute rottmdatis (nee emarginatis) ad bases abrupte 
constrictis (nee rectis), secus margines crenatis (nee integerrimis) confeste separabilis. 

Terrestrial dioecious tree to 8 m tall. Branchlets angulate, 8—10 mm diam., 
densely and minutely rubiginous furfuraceous-lepidote, conspicuously ru- 
biginous punctate-lineate below. Leaves pseudoverticillate; blades coriaceous, 
oblanceolate, (21— )26— 34.5 cm long, (6.5-)9— 12.2 cm wide, apically acute 
to subacuminate, basally acute, decurrent on the petiole 4—7 mm, glabrous 
above, very minutely rubiginous furfuraceous-lepidote below, the midrib 
slightly raised above, prominently raised and rubiginous punctate-lineate 
below, the secondary veins 7-10 pairs, slightly impressed above, promi- 
nently raised below, the margin entire, flat; petioles canaliculate, 2.5—3 cm 
long, swollen below to 0.5—0.7 cm diam. basally, rubiginous furfuraceous- 
lepidote at first, early glabrescent, conspicuously rubiginous punctate-lineate. 
Staminate inflorescence: a bipinnate panicle (12— )14— 17.5(— 25) cm long, 2—3 
cm wide; peduncle (l-)2— 3.5 cm long; secondary inflorescence bracts chartaceous, 
linear, 2—3 mm long, 0.3—0.6 mm wide, apically attenuate, densely glan- 
dular-papillate, the margin entire; branches racemose-glomerulate (0.5— )1 — 
1.5 cm long; floral bracts membranaceous, linear, 1.6—1.8 mm long, 0.3— 
0.4 mm wide, apically attenuate, densely rubiginous puberulent; pedicels 
cylindrical, 0.3—0.5 mm long, glabrescent. Staminate flowers 4-merous, pink; 
calyx membranaceous, cotyliform, 1.4—1.6 mm long, the tube ca. 0.2 mm 
long, the lobes ovate, 1.2—1.4 mm long, 1—1.1 mm wide, apically acute, 
sparsely and inconspicuously orange punctate, glabrous, the margin coarsely 
crenulate, glabrous; corolla membranaceous, subrotate, 2.3—2.5 mm long, 
the tube 0.2—0.3 mm long, densely glandular-granulose, the lobes ovate, 
2-2.3 mm long, 1 .7—1 .8 mm wide, apically acutely rounded, abruptly constricted 
basally, inconspicuously orange punctate without, sparsely gladular-granulose 
behind the base of the filaments, the margin hyaline, coarsely crenate, gla- 
brous; stamens 1.8—2 mm long, the tube carnose, squarrose, 0.2—0.3 mm 
long, the filaments terete, 1.1—1.2 mm long, slightly curved proximally, 
the anthers ovate, 0.5—0.6 mm long, 0.6—0.7 mm wide, apically rounded, 
basally cordate, the connectives inconspicuously orange punctate dorsally; 
pistillode obturbinate, 0.5 mm long, 0.6 mm diam., the stigma truncate, 
minutely lobed. Pistillate inflorescence as in staminate but (3.5-)5.5-9-5 cm 
long, the branches glomerulate. Pistillate flowers as in staminate but red, 
except white on corolla lobe apices; calyx obconic, 1.3—1-7 mm long, the 
tube 0.6—0.8 mm long, the lobes very widely ovate to suborbicular, 0.7— 

106 SiDA 18(1) 

1.2 mm long, 0.8—1.2 mm wide, apically broadly rounded, corolla 2.3—2.7 
mm long, rhc cube ().6-{).8 mm long, the lobes suborbicular, 1.5—1.7 mm 
long, 1.5—1.8 mm wide, the margin irregtilar, hyaline, staminodial tube 
1-1.3 mm long, the apically free portions of the filaments 0.4-0.5 mm long, 
the antherodes 0.3—0.4 mm long and wide; pistil obnapiform, 2.2—2.4 mm 
long, 1.3—1.5 mm diam, the ovary 1.3—1.5 mm long, the style thick, 1.1— 

1 .3 mm long, the stigma capitate, 4-lobed, the lobes recurved, the placenta 
deeply cuptililorm, the ovtdes 2—3, imbedded. Fr//it unknown. 

Distribution. — Known only fi'om the Department of Loreto, in Alto Amazonas 
and Loreto Provinces, Peru, at 160—220 m elevation. 

Ecology and amservci turn stc/tns, — Cybianthiis I'c/sqi/ezii occurs in primary terra 
jirnie lowland forest, with scattered white sand areas. Label data do not al- 
low determination of whether this species occurs on the laterite or on the 
sands. Because it is known only from two gatherings, its conservation sta- 
tus is unknown. However, its importance as a tonic in Mayna Jivaro cul- 
ture may indicate it is locally common. 

Etymology. — It gives me great pleasure to dedicate this species to a great 
friend and colleague, Ing. Rodolfo Vasquez Martinez, Assistant Curator and 
Director of the Flora of Peru Program of the Missouri Botanical Garden. 
Vasquez is an indefatigable collector, a forestry engineer, dendrologist, taxonomist 
and author of numerous publications on uses of Peruvian forest products, 
economic plants of the Peruvian Amazon, and Flon/la of the Biological Re- 
serves of hinitos. He is a taxonomic authority on Caraipa and Myristicaceae 
of the Amazon Basin. 

Local names and uses. — Peru: "sesa," "kurup" (Mayna Jfvaro). The sap is 
extracted and a juice is drunk to "improve hunting ability." 

PARATYPii: PERU. Loreto: Prov. Loreto, Pampa Hermosa and vicinity, Rio Corrientes, 1 
km S of junction with Ri'o Macusan, 03° 15' S, 75° 50' W, 160 m, 3-20 Dec 1985 (pist. 
fl), W. Lcins et at. 10306 (BRIT, MO). 

Cybianthits vascyiezii is most closely related to C potiaei c^f the eastern Amazon 
Basin (French Gtiiana and Brazil (Amapa, Bahfa), but is easily recognized 
by the conspicuously rubiginous punctate-lineate branchlets, larger 
pseudoverticillate, oblanceolate leaves, longer petioles and inflorescences, 
ovate, apically rounded corolla lobes that are abruptly constricted basal ly, 
and coarsely crenate along the margins. 

26. Cybianthus cenepensis Pipoly, sp. nov. (Fig. 20). Tyi^i : PERU. Amazonas: 
Ri'o Cenepa, vicinity Huampami, ca. 5 km E of (Chavez Valdi'via, 04° 30' S, 78° 30 
W, 200-250 m, 1 2 Aug 1978 (stam. fl),/\, K/ijikcit 263 (iiolotyph: MO; isotypi.s: F, 
MO, NY, US, USM). 

Ob folia chartacea oblanceolata atl apices acummara ad bases ciineata, petiolos canaiicnatos, 
inHorescentias angnste bipinnatipaniciilatas, ad C. h/icbtieiii valde atfinis sed ab ea ramnlis 
alatis (nee laevibtis), pctiolis 2-2.5 (non 1 .5—1 .8) cm longis, lobis calycinis late ovatis (ncc 

PiPOi.'i', Cybianthus in Ecuador and PerLi 



Fig. 20. Cyhianth/ts cmepensis Pipoly. A. Habit, showing winged branchlers. B, Sraminace 
inflorescence branch, showing secondary inflorescence branch bracts. C. Staminate flower, 
showing stamens subeqtial to corolla lobes, conspicuous square staminal tube, and coarsely 
crenate corolla lobes. D. Pistillate flower, showing ellipsoid pistil. E. Pistillate inflorescence 
branch, showing dense spike appearing glomerulate. D. Branchlet apex, showing puberu- 
lent vestiture. A drawn from holotype. B— D, drawn from Auc/uish 522. E— F, drawn from 
Kujikat 306. Figure drawn by Linda Ellis. 

108 SiDA 18(1) 

lineari-lanccolatis), grossc crenaris (ncc inte^'errimis), lobis corollinis grosse crcnatis (nee 
enccris) clenicjue antlieris ad a|-iices acLitis (nee roninclatis) atl bases eordatis (nee obtLisis) 
perhieile clisringuitLir. 

Treelet to 2 m tall. Branchkts prominently longitudinally ridged, the ridges 
forming small, rounded wings, (2.5-3-)5-6 mm diam., sparsely rufous 
puberulent, glabrescent. Leaves alternate; blades chartaceous, oblanceolate, 
(1 3.7— )18.5-23(— 26.2) cm long, (4— )6-10 cm wide, apically acuminate, 
basally cuneate, glabrous above, sparsely rufous puberulent below, glabres- 
cent, inconspicuously pellucid punctate, the margin entire, flat; petioles 
canaliculate, 2—2.3 cm long, sparsely rufous puberulent, glabrescent. Staviinate 
inflorescence a lax bipinnate panicle, 11—1 8 cm long, 1—1.7 cm wide, the rachis 
densely rufous papillate; secondary inflorescence bracts chartaceous, linear- 
lanceolate, 4—4.5 mm long, 1—1.2 mm wide, apically subulate, densely rufous 
tomentellotis, the margin irregular, entire; inflorescence branches 3-8 mm 
long, the flowers densely subspicate, appearing glomerulate; floral bracts 
chartaceous, linear, 1 — 1.2 mm long, 0.1— 0.2 mm wide, apically subulate, 
densely glandular-papillate, the margin crenulate, glabrous; pedicels cylindrical, 
0.2—0.3 mm long, densely glandular-papillate. Staminate flowers A-mtrons, 
chartaceous, brownish-purple, 1.6—1.8 mm long; calyx cotyliform, 1—1.2 
mm long, the ttibe 0.3—0.4 mm long, the lobes broadly ovate, 0.7— 0.8 mm 
long, 0.6—0.7 mm wide, apically acute, prominently brown punctate, the 
margin hyaline, coarsely crenate, glabrous; corolla subrotate, 1.2—1.3 mm 
long, the tube ca. 0.1 mm long, densely glanclular-granulose within, the 
lobes widely ovate, 1 — 1 .2 mm long, 0.8—1 mm wide, apically subacuminate, 
glabrous and inconspicuously orange punctate without, glabrous within except 
under the filaments, the margin hyaline, coarsely crenate, glabrous; stamens 
subequalling the corolla lobes, 1 — 1.1 mm long, the tube conspicuous, carnose, 
square, 0.1-0.2 mm long, densely glandular-granulose, the filaments 0.6— 
0.7 mm long, the anthers ovate, 0.3—0.4 mm long, 0.4—0.5 mm wide, apically 
acute, basally cordate, the connective inconspicuously orange punctate; pistillode 
obturbinate, 0.1—0.2 mm long, 0.2—0.3 mm diam., hollow, glabrous. Pis- 
tillate inflorescence diS in staminate but 12—14 cm long, 0.8—1 .0 cm wide; secondary 
inflorescence bracts 3.8—4. 1 mm long, 0.8—1 mm wide; inflorescence branches 
3.5—6 mm long; floral bracts 1 — 1.2 mm long, 0.1—0.2 mm wide; pedicels 
0.2—0.3 mm long. Pistillate flowers as in staminate but white, 1.1 — 1.4 mm 
long; calyx 0.6—0.8 mm long, the tube 0.1—0.2 mm long, the lobes 0.5— 
0.6 mm long, 0.3—0.4 mm wide; corolla 0.7—0.8 mm long, the tube ca. 
0.1 mm long, the lobes 0.6— 0.7 mm long, 0,5—0.6 mm wide; staminodes 
much sorter than the corolla lobes, 0.3—0.4 mm long, the tube conspicu- 
ous, carnose, circular, ca. 0. 1 mm long, the filaments ca. 0.1 mm long, the 
anthers 0.2—0.3 mm long, 0. 1—0.2 mm wide; pistil ellipsoid, 0.5—0.6 mm 
long, 0.2—0.3 mm diam., the style 4-lobed, the lobes curved distally, pla- 

PiFOLY, Cybianthus in Ecuador and Peru 109 

ccnta subobose, bearing 4 partially immersed ovules. Fruit globose, 6—7 
mm long and in diam., prominently red punctate. 

Distribution. — Known only from the Rfo Cenepa Drainage Basin, Amazonas, 
Peru, 200—250 m elevation, and from one disjimct population from Napo, 
Ecuador, very close to the Colombian border. 

Ecology and conservation status. — Cybianthus cenepensts inhabits varzea for- 
ests in the Amazon of Ecuador and Peru. Despite lack of collections, it is 
known that the Napo and the Cenepa Rivers are shallower than many with 
their same volume, thus giving them strong currents. Aside from the need 
for much more exploration at the edge of the Amazon Basin in Ecuador and 
Peru, perhaps the strong current of these rivers makes the dynamics of their 
adjacent varzea forests different from others in Amazonia. At this time, the 
conservation status of this species is unknown. 

Etymology. — The specific epithet refers to the region of Peru from which 
the type was collected, the Rio Cenepa Drainage Basin of the Alto Amazonas 
Province, Amazonas Department, Peru. 

Local names and uses. — Peru: "sauka" (Huambisa). 

Paratypes. ECUADOR. Napo: Canton Orellana, Yasuni National Park, Maxus Rd. and 
pipeline construction project, km 15. 01° 3 I' S, 76° 32' W, 250 m, 30Jun 1994 (bud), N. 
Pitman 461 (BRIT, MO, QCNE); Rio Yasunf, periodically inundated forest ca. 80 km upriver 
from Nuevo Rocaluerte, 225 m, 17 Sep 1 977 (ft), R. Foster 3708 (F, QCA). PERU. Amazonas: 
Quebrada Sasa, Rio Cenepa, 250 m, 2 Jun 1973 (stam. fl), E. Ancuasb 522 (MO, USM); 
Ri'o Cenepa, vicinity of Huampami, ca. 5 km E of Chavez Valdivia, ca. 4° 30' S, 78° 30' W, 
200-250 m, 12 Aug 1978 (pist. fl), A Kupkat 306 (F, MO, USM); Quebrada Chigki Shiunk, 
4° 30' S, 78° 30' W, 11 Aug 1978 (bud), £, Amtiash 1412 (MO, USM). 

Cybianthus cenepensis is closely related to C. buchtieni Pax of the Mapiri 
region in Bolivia. However, Cybianthus ce?iepensis is easily recognized by its 
winged branchlets, petioles 2—2.5 cm long, widely ovate and coarsely crenate 
calyx lobes, coarsely crenate corolla lobes, and anthers with acute apices and 
cordate bases. The ellipsoid pistil is also unique within the subgenus. 

27. Cybianthus nanayensis (J.F. Macbr.) G. Agostini, Acta Biol. Venez. 
10:l60. 1980. Weigeltia nanayensis ].V. Macbr., Field Mus. Nat. Hist., Bot. Set. 
11:33. 1 931. Type: PERU. LORETO: Lower Rfo Nanay, 4 Jun 1929 (stam. fl),L/. Williams 
658 (holotype: F; fragment, G). 

Weigeltia silvestris ].¥. Macbr., Candollea 6; 16. 1934. syn. nov. Cybianthus silfestris (J.F. 
Macbr.) G. Agostini, Acta. Biol. Venez. 10:163. 1980. Type: PERU. Loreto: Mishuyacu, 
near Iquitos, Dec 1929 (stam. fl), G. King 724 (holotype: F; isotypes: NY, US). 

Cononiorpha d/ibia } .V. Macbr., Candollea 6: 17. 1934. syn. nov. Cybianthus dubius (^ .V. Macbr.) 
G. Agostini, Acta Biol. Venez. 10: 158. 1980. Type: PERU. Loreto: Mishuyacu, near 
Iquitos, 24-28 Sep 1929 (pist. fl, fr), A. Killili & A. Smith 29906 (holotype: F; isotypes: 
NY, US) [erroneously cited as G. King 299(^6]. 

Conomorpha loretensis Lundell, Wrightia 6:113. 1 980. syn. nov. Cybianthus loretensis (Lundell) 
Pipoly, Brittonia 33:496. 1981. Type: PERU. Loreto: Quistococha, near Iquitos, 18 
Nov 1977 (stam. fl), A. Gentry 20763 (holotype: LL-TEX; isotypes: F, MO, NY). 

1 10 SiDA 18(1) 

S//hshr//h to shrub to ().5(— 1.5) m tall. Braiichlets 0.2—0.5 cm thick, smooth, 
rufoLis-lepidote. Leai'es in loose pseudoverticels; blades chartaceous to sub- 
coriaceous, elliptic, lanceolate or oblanceolate, (7— )1 l-18(-25.5) cm long, 
(2.9~)3.5— 7.5(— 10) cm, apically acute to acuminate, basally acute to cu- 
neate, midrib depressed above, prominently raised below, the secondary veins 
7—16 pairs, glabrous above, sparsely and minutely rutous-lepidote below, 
puncticulate, the margin flat, entire; petioles marginate, (0.6— )().9— 1.5 cm 
long, glabrous. Inflorescences monomorphic, a simple raceme or rarely with a 
second, malformed basal branch, (1 .5— )3— 9-5(— 14.5) cm long, the rachis 
densely glandular-papillate; floral bracts membranaceous, linear-lanceolate, 
0.2—1.2 mm long, 0. 1—0.2 mm wide, the apically long-attenuate, sparsely 
glandular-papillate, the margin glandular-ciliate, entire; pedicels cylindri- 
cal, (0.7— )1— 1 .5(— 2) mm long, densely glandular-papillate. Stan/inate flowers: 
white to cream, 4-merous, chartaceous; calyx cotyliform, 1—1.3 mm long, 
the tube ca. 0.1 mm long, the lobes widely ovate to deltate, 0.9—1.2 mm 
long, 0.8—1.2 mm wide, apically acute to actnninate, medially thickened 
and prominently black punctate, glabrous, the margin hyaline, irregular, 
densely glandular-ciliolate; corolla rotate, 1.9—2.3 mm long, the tube 0.6— 
0.8 mm long, the lobes widely ovate to ovate, 1.3—1-6 mm long, 1.2—1.4 
mm wide, apically acute to rounded, subentire or apically notched, medi- 
ally thickened and prominently black punctate, glabrous without, glandu- 
lar-granulose within at junction of tube and lobe, the margin hyaline, ir- 
regular, glabrous; stamens 2.2—2.5 mm long, exserted or rarely subequal to 
corolla, the staminal tube carnose, conspicuous, 0.5—0.7 mm long, subtruncate 
between the filaments, the apically free portions of the filaments terete, (1 .2—) 
1.4—2 mm long, the anthers widely ovate, 0.32-0.36 mm long, apically 
rotmded to obttise, basally cordulate, the connective prominently punctate 
dorsally; pistillode vestigial, obclavate or tubiform, 0.7—0.8 mm long, sparsely 
translucent glandular-lepidote, hollow. Pistillate flowers as in staminate but 
beige to brown; calyx 2—2. 1 mm long, the tube 0.1—0.2 mm long, the lobes 
1.9—2 mm long, 1.9—2 mm wide; corolla cotyliform, 1.9-2 mm long, the 
tube 0.5—0.6 mm long, the lobes widely ovate, 1.2—1.4 mm long, 1.3—1.4 
mm wide, apically acute to acuminate; staminodes resembling stamens, 1.2— 
1.3 mm long, the staminodial tube 0.5—0.6 mm long, the filaments 0.4— 
0.5 mm long, the antherodes subquadrate, ca. 0.2 mm long and wide, apically 
obtuse, basally truncate; pistil ellipsoid, 1.4—1.6 mm long, the ovary 1 — 
1.2 mm long, 1.1—1.2 mm diam., densely translucent glandular-lepidote, 
the placenta cupuliform, ovules 2, partially immersed, the style trunctae, 
0.3—0.4 mm long, the stigma punctiform. Fr//it depressed-globose, 4.5— 
5.5 mm long, 5.5— 6.5(— 7) mm diam., yellow when fresh, prominently black 

PiPOi.Y, Cybianthus in Ecuador and Peru 111 

Distribution. — Endemic to the tall moist forests on white sands of the 
Peruvian Amazon, primarily from the Iquitos area, 100—160 m. 

Ecology and conservation status. — Cybianthus nanayensis is locally common, 
and thrives in gaps left by large treefalls in overmature forests, and along 
the margins of forest margins and paths, where it occurs in a rather dense 
herbaceous layer. However, it does not tolerate compacted soils. With de- 
creasing quantitites of habitat owing to logging pressures, the species should 
be considered threatened. As a gap species growing on nutrient deficient 
soils, and with very attractive fruits, Cybianthus nanayensis shows great promise 
as a potentially marketable horticultural plant. 

Etymology. — The epithet takes its name from the river basin where it occurs, 
the Nanay River. 

Representative specimens examined. PERU. Loreto: Prov. Loreco, Nauta, 0-4°32' S, 73°35' 
W, 160 m, 2 Jun 1984 (stam. fl), R. Vdsquez & N.Jaramillo 3073 (AMAZ, MO, NY), (fr), 
R. Vasquez & N.Jaramitlo 3086 (AMAZ, MO); Prov. Maynas, Allpahuayo, IlAP Experi- 
mental Station, 04° 10' S, 73° 30' W, 120 m, 20 Sep 1990 (fr),J. Pipolyetal. 12263 (AMAZ, 
MO, USM), 15 Aug 1990 (fr), R. Vasquez & N.Jaramillo 14204 (AMAZ, MO, US, USM), 
1 Oct 1990 (stam. fl), R. Vasquez & N.Jaramillo 14463 (AMAZ, MO); Laguna Quistococha, 
1 5 km SW of Iquitos, 8 Jul 1977 (stam. fl),J. Solomon 3466 (LL-TEX, MO); Mjshiana, 30 
km SW of Iquitos, Callicebus Biological Reserve, Ri'o Nanay, 4 km S of Mishana, 19 Aug 
1978 (stam. fl), R. Foster 4243 (AMAZ, F), 16 Aug ] 980 (stam. fl), 4327 (F-2 sheets); Vicinity 
Mishana, between Rfo Nanay and Rfo Itaya, 130 m, 29 Nov 1977 (fr), A. Gentry et al. 
21033 (F, MO); Between Iquitos and Sta. Maria de Nanay, 180 m, 31 May 1978 (stam. fl), 
A. Gentry et al. 22367 (AMAZ, F, MO); 03° 50' S, 73° 30' W, 25 Feb 1981 (fr), A. Gentry 
et el. 31479 (AMAZ, MO), 31 Dec. 1 982 (mixed- stam. fl, fr), A. Gentry & L. Emmons 38776 
(MO-2 sheets), 5 Aug 1990 (fr), R. Vasquez el al. 14161 (AMAZ, MO, USM); Rfo Nanay, 
May-Jun 1929 (stam. fl), LI. Williams 637 (F); Mishuyacu, near Iquitos, 100 m, Oct-Nov 
1929 (stam. fl), G. Klug 304 (F), May-Jun 1930 (stam. fl), G. Klug 1335 (F); 6 Oct. 1982 
(stam. bud), R. Vasquez & N.Jaramillo 3261 (MO), 20 Jan 1985 (stam. fl), 6123 (MO); 
Puerto. Almendras, 03° 48' S, 73° 25' W, 122 m, 4 Jan. 1986 (pist. fl), R. Vasquez & N. 
Jaramdlo 7070 (AMAZ, MO), 30 May (stam. bud), 7393 (MO), 20 Oct. 1986 (fr), 8073 
(AMAZ, MO); Roca Fuerte (Momon), Oct-Nov 1984 (stam. fl), R. Vasquez & N.Jaramillo 
3241 ■ Madre de Dios: Prov. Tambopata, Tambopata Reserve, Rfo Tambopata at mouth of 
Rfo D'Orbigny, 250 m, 6 Mar 1981 (fr), A. Gentry & K. Young 32023 (AMAZ, MO, NY). 

Agostini (1980) had not seen the type specimen of Weigeltia silvestris and 
included it in Cybianthus subgenus Comomyrsine, probably due to matching 
with a herbarium misidentification of a specimen of Cybianthus kayapii (Lundell) 
Pipoly as Weigeltia silvestris in NY. I previously recognized Cybianthus dubius 
and C. loretensis as distinct taxa (Pipoly 1981) on the basis of quantiative 
characters, and had not seen the type of C silvestris. The exserted stamens 
and obclavate or tubiform pistillode of the staminate flowers, and truncate 
style with punctiform stigma in the pistillate flowers are unique features 
within the subgenus. 

112 SiDA 18(1) 

Populations corresponding to the type oiWeigeltia silvestris have sHghtly 
larger leaves, but are otherwise indistinguishable from the type of Wejgeltia 
nanayemh. The type oi Conomorpha di/bia is notable only for differences at- 
tributable to the fact that it is a pistillate fruiting specimen. Fieldwork in 
Peru has shown that populations corresponding to the type oi Conomorpha 
loretensis grow in full sun, and consequently have narrower leaves with longer 
petioles than those of the type of Weigeltia nanayensu . 

VII. Cybianthus subgenus Grammadenia (Benth.) Pipoly, Mem. New York 
Bot. Gard. 43:47. 1987. GrammackmaV>ftnx.\\., PI. Hartw. 218. 1846. Type Spi-riES: 
Grannuadenui niarginata Benth. = Cyhiaiithus iiiarginati/s (Benth.) Pipoly. 

Erect or pendent, evergreen, terrestrial, epiphytic or epipetric dioecious, 
bisexual, dioecious or monoecious shn/bs or small trees. Roots diageotropic. 
Bark mostly smooth, gray or sometimes brown, cracking transversely on 
older parts. Trunks distinguishable, normally more or less terete, leptocaulous 
or rarely pachycaulous basally, growth dynamics corresponding to Rauh's 
Model (Halle et al. 1978). Branchlets thin to moderately thick, terete or ridged, 
smooth to verruculose to verrucose, glabrous or rarely glandular-papillate 
apically, glandular-papillate at first in the leaf axils then glabrescent, with- 
out lenticels. Cataphylls and pseudocataphylls absent. Leaves alternate, 
supervolute, exstipulate, sessile, acrodromous, apically obtuse-mucronate, 
basally auriculate, subamplexicaul, minutely glandular-papillate along midrib 
above, glabrescent, glabrous below, hydropotes few or absent above, nu- 
merous below, the margins hyaline, membranaceous, prominently punc- 
tate and punctate-lineate, entire or minutely crenulate to denticulate; peti- 
oles absent. Inflorescence monomorphic, simple, axillary raceme, at times reduced 
to appear dichasial; inflorescence bract broadly ovate to deltate, acute, prominently 
punctate, the margins entire to erose, early caducous; rachis terete, straight 
or flexuous, minutely glandular-papillate, glabrescent; floral bracts solitary, 
at pedicel base, ovate to lanceolate, persistent; pedicels cylindrical to clav- 
ate, minutely glandular-papillate, glabrescent, accrescent in fruit. Floivers 
unisexual or bisexual, (4— )5— 6(— 7)-merous; perianth lobes imbricate or 
quincuncial or rarely and aberrantly dextrorsely contorted, promiently, con- 
spicuously or inconspicuously punctate and punctate-lineate; calyx cotyliform, 
the lobes erect, at times reflexed in fruit, the margins irregular or regular, 
entire or minutely crenulate, glabrous or glandular-ciliate; corolla rotate, 
bearing a ring of glandular granules at the junction of tube and lobe, the 
lobes glabrous without, glabrous or rarely glanduiar-granulose and smooth 
or rugose within, the margins regular or irregular, entire or minutely crenulate; 
stamens and staminodes similar, connate to form a conspicuous, membranaceous 
or carnose tube adnate to the corolla tube, elobate or with minute lobes 
alternating with the anthers, the anthers and antherodes basifixed, sessile 

PiPOLY, Cybianthus in Ecuador and Peru 113 

or on minute apically free filaments, quadrate or ovate, apically emarginate 
or rounded, basally truncate, dehiscent by apical birimose pores, usually 
dorsally and rarely ventrally punctate; pistil and pistillode similar, obnapiform, 
ellipsoid or umbonate, the ovary terete, lobed or costate, sparsely to densely 
translucent-lepidote, the placenta umbonate, (l-)2-3(-4) ovulate, the ovules 
half-immersed in the basal placenta, uniseriate or biseriate, the style short, 
truncate, the stigma punctiform. Fn/it drupaceous, depressed globose, obovoid, 
or ellipsoid, the exocarp sometimes fleshy, the mesocarp and endocarp stony, 
prominently punctate-lineate, white, lavender or purple-black at maturity, 
one seeded, the testa corrugate, the embryo cylindrical, transverse. 

A subgenus of 7 species, in the Lesser Antilles, in Mesoamerica from northern 
Costa Rica through Panama, in the Andes from Venezuela to southern Peru, 
east through the Guayana Highland and to the Serrania de Turumiquire, 
Anzoategui, Venezuela. In Ecuador, two species occur. In Peru, 3 species 
have been recorded, of which one, Cybianthus lineatus (Benth.) Pipoly, for- 
merly thought to be a Guayana Highland endemic (Pipoly 1987) is reported 
for the first time here. 


1 . Branchlets verruculose- to verrucose-papillate or red glandular-papillate; leaves 

coriaceous, symmetrical; inflorescence erect, flexuous; perianth coriaceous; 

epipetric or terrestrial shrubs or trees ot lagunas in paramos and elfin forests 

or in "jalca" or "pajonal" alpine savannas. 

2. Epipetric shrub or tree to 6 m tall; trunk leptocaulous; branchlets verruculose- 
to verrucose-papillate, prominently ridged, rugose when dried; leaves oblong, 
elliptic or lanceolate, (l.l-)1.4-2(-2,5)cm wide; fruit ovoid, then ellip- 
soid at maturity; plants of paramos or elfin forests 28. C. marginatus 

2. Terrestrial shrub to 1.5 m tall; trunk pachycaulous; branchlets red glan- 
dular-papillate, terete, smooth when dried; leaves oblanceolate, (0.4-)0.6- 
1(-1.2) cm wide; fruit obovoid throughout development; plants of jalca 

or pajonal alpine savannas 29- C. lineatus 

1 . Branchlets smooth, glabrous; leaves chartaceous, asymmetrical; infiorescence 
lax, straight; perianth chartaceous; epiphytic shrub or tree in cloud lorests 
below subparamos or facultative epiphytes in montane and elfin forests. 
30. C. magnus 

3. Leaves not bearing hydropoten above, conspicuously black punctate and 
punctate-lineate below; inflorescence rachis black punctate-lineate; peri- 
anth whitish-green, prominently black punctate and punctate-lineate; anthers 
ovate, rounded apically; branchlets 4-7 mm diam.; fruit purple-black at 
maturity 30a. C. magnus subsp. magnus 

3 . Leaves bearing hydropoten above, at least proximally, inconspicuously pellucid 
to orange punctate; inflorescence rachis orange punctate-lineate; perianth 
maroon to purple, rarely white (then pistillate), prominently orange-punctate; 
anthers quadrate, emarginate apically; branchlets 2. 5-3. 5(— 4.0) mm diam.; 
fruit white, then lavender at maturity 30b. C. magnus subsp. asymmetricus 


SiDA 18(1) 

Fig. 21. Piccoriaiized distribution of C. marginatm. A-K, Variation in leaf shape, marginal 
venation and punctation; note prominent apical mucron, sessile leaf base typical of subge- 
nus Grammadema. A-K, drawn from: A. Cuatrecasas 21803, B. Pipoiy 6954, C. Pipoiy 6539, 
D. LmiuT 8038, E. L/iteyn 9032, F. Steyermark 100867, G. Mason 13730, H. Pfpoiy 6975 , 
I. Pearce 250, ]. Leh//unin 399, K. Luteyn /Oi 7 J. Figure from Pipoly, 1 987, drawn by Bobbi 

28. Cybianthus marginatus (Bench.) Pipoly, (Fig. 1 A,B, 7C,F, 21). Mem. 
New York Bot. Gard. 43:60. 1987. Grammadema marginata^mxh., PI. Hartw. 
218. 1846. Type: COLOMBIA. Cauca: Near Pitayo, 3,636 m, 1843 (bisex. fl), C. 
Uartweg 1200 (holotype: K; isotypes: BM, E, G-BOISS, G-DEL, LD, OXF, P, W-2 


Grammadeu/a lehmaniiii Mez in Engl., Pflanzenr. IV. 236(Heft 9): 231. 1902. Type: 
COLOMBIA. Toljma: Altos de Otesas, 3,300 m, 1 1 Jan 1883, (bisex. fl), F. Lehmann 
2399 (ei-CTOTYPE by Pipoly 1987: G; isolectotypes: LE, US). 

PiPOLY, Cybianthus in Ecuador and Peru 1 15 

Grcnumackma alp'ma Mez in Engl., Pflanzenr. IV. 236(Heft 9):231. 1902. Typh: VEN- 
EZUELA. Andes of Trujillo and Merida, 1,212-4,390 m, 1842 (bisex. fl),J. Linden 
447 (lectotype by Pipoly 1987; P; isoleci-otypes: BM-2 sheets, BR, G, G-DEL, G- 

Grainmadenia pastensis Mez in Engl., Pllanzenr. IV. 236(HeFt 9):232. 1902. Type: CO- 
LOMBIA. Narixo: W Cordillera of Pasto, 3, 000-3, 200 m, 20 Feb 1881 (bisex. fl), F. 
Lehmann 399 (holotype: G; jsotypes: BM, LE). 

Grammadema mttda Mez in Engl., Pflanzenr. IV. 236(Heft 9):232. 1902. Type: PERU. 
Hl;Anuc:o: Pozuzo, 2,131-2,727 m, 1863 (bisex. fl, fr), R. Pearce 250 (holotype: K). 

Grammadema weberbaueri Mez, Repcrt. Spec. Nov. Rcgni Veg. 1 6:418. 1920. Type: PERU. 
Cajamarca: Jaen, cordillera E of Huancabamba, E slopes, 2,400-2,500 m, Apr 1 91 2 
(bisex. fl), A. Weberbauer 6121 (lectotype by Pipoly 1987: GH; isolectotype: F). 

Grammmadmia hexamera Pittier, J. Wash. Acad. Sci. 21:l40. 1931. Type: VENEZUELA. 
Merida: Tabay, 2,500-3,000 m, 18 Sep 1930 (bisex. fl), W. Gehriger 47 1 (holotype: 
VEN; isotypes: A, F, G, NY, PH). 

Grammadenia andkola Cuatrec, Revisca Acad. Colomb. Ci. Exact. 8(3 1):32 1 . 1 95 1 . Type: 
COLOMBIA. Valle: Cordillera Occidental, Los Farallones, NW slope, Qucbrada Las 
Nieves, below El Diamante, 2,900 m, 30 Jtil 1 946 (bisex. fl, fr),/. Ciuitrecasas 21805 
(lectotype by Pipoly 1987: F; isoLE.cTcrrYPE.s: F, COL 2-sheets, U, US). 

Epiphytic shrub or small tree to 6 m, the trunk leptocaulous. Branchlets 
prominently ridged, 3-4(-5) mm diam., verruculose- to verrucose-papil- 
iate, rugose when dried, glabrous. Leaves coriaceous, symmetrical, oblong, 
elliptic or lanceolate, (3-)3.5-6(-6.5) cm long, (l.l-)1.4-2(-2.5) cm wide, 
apically acute to obtuse, mucronulate, basally acute, auriculate, nitid above, 
pallid below, prominently punctate and punctate-lineate, the margin en- 
tire, revolute. Inflorescence erect, the rachis flexuous, slender, (1— )1.3-2(— 3.2) 
cm long, densely black punctate-lineate; floral bracts widely ovate to deltate, 
(().8-)1.2-1.5(-1.8) mm long, 0.8-1.4(-I.8) mm wide, apically acute, the 
margin entire; pedicels (1-)1 .5-2 mm long. Flowers coriaceous, 5(-7)-merous; 
calyx 1.1-2.1 mm long, the tube ca. 0.1 mm long, the lobes deltate, (1-) 
1.5-2 mm long, (1.2-)1.3-1 .5(-2) mm long, apically acute, prominently 
and densely punctate and punctate-lineate, the margins minutely crenu- 
late, glabrous to minutely ciliolate at first apically; corolla (2-)2.2-3 mm 
long, the lobes widely ovate, (l-)1.2-2 mm long, (1.2-)1.5-2(-2.5) mm 
wide, obtuse to rounded, glabrous without, glandular-granulose within basally, 
densely and prominently punctate medially, the margin entire; staminal tube 
carnose, 0.6-0.8(-l) mm long, the anthers sessile, alternate with fleshy lobes 
to 0.2 mm long, quadrate, 0.4-0.7 mm long and wide, apically rounded, 
basally truncate, the connective prominently punctate dorsally; pistil obnapiform, 
(0.9-)1.2-1.5 mm long and (0.8-)1.2-1.6 mm diam., the ovary (0.4-)0.6- 
1 mm long, glabrous to sparingly translucen glandular-lepidote, ovules 2(- 
4), uni- or biseriate, the style 0.3-0.5 mm long, epunctate. Frai/ somewhat 
ovoid, then ellipsoid at maturity, 4.5-5(-6) mm long, (2.5-)3-4 mm diam., 
white, then purple-black at maturity, the punctations red-black, prominent. 

Distribution. — Cybianthus marginatus is the most common species of the 

116 SlDA 18(1) 

subgenus, occurring throughout the Andes from Venezuela to Peru, 2,000- 
3,400 m. 

Ecology and anuervatum it at US. — This species occurs in large populations 
on rocks above lagunas in paramos in northern Ecuador and along water- 
courses in paramoid elfin forests, and "ceja" formations in the remainder of 
Ecuador and Peru. As long as there are remnant paramo formations, or elfin 
forests, there will be populations o^ Cyhianthus marginatm. However, in some 
instances, soil compaction due to overgrazing by sheep can render the soil 
uninhabitable for this species. 

Etymology. — The specific epithet refers to the scarious leaf margin, made 
conspicous by the coriaceous texture. In addition, this aspect is made more 
conspicuous by the prominent submarginal vein of many populations. 

Representative specimens examined. ECUADOR. Azuay: "Oriente" border. Paramo deJ 
Castillo, crest of E cordillera on trail between SeviUa de Ore and Mendez, 2,727-3,333 m, 
18 Aug 1945 (fl, fr), W. Camp E-4809 (NY, VEN); Rio Collay, Huagarancha S of El Pan, 
2,650-3,290 m, 6 Jtd 1 943 (fl) J. Steyermark 53354 (NY), carchi: Peak of Cerro Golondrinas, 
00° 51' 12" N, 78° 08' 2 1" W, 3070 m, 24 Jul 1994 (fr), B. Boyle et at. 3373 (BRIT, MO, 
QCNE); Canton Montufar, Loma El Corazon, Bretaiia, SE of Mariscal Sucre, Ri'o Minas, 
00° 35' N, 77° 42' W, 3,150 m, 22-23 Dec 1992 (fl, fr), W. Palacios & G. Tipaz 10569 
(BRIT, MO, QCNE). Imbabura: Cordillera Oriental, Camp Arelan, E of Volcan Cayambe, 
2,803 m, 21 Jul 1944 (fl, fr), W. Drew E-351 (MSC); Ridge just S of Rio Clavadero, along 
trail to Rio San Pedro, E of Cayambe, 2,893 m, 27 Jul 1944 (fl, fr), /, Wii^gn/s 10484 (DS, 
US). Loja: Saraguro-Loja, Km 12.4, turnoff toward Fierro Urco, Km 2.5-2,7, 03° 41' 05" 
S, 79° 17' 20" W, 3,150-3,300 m, 7 Dec 1994 (fl), P.Jorgensen et at. 1278 (BRIT, LOJA, 
MO, QCA, QCNE); Paramos de Saraguro, 10 km S of Saraguro, 3,050 m, 2 Jan 1979 (fl), 
_/. Luteyn et al. 6647 (NY, QCA). Zamora-Chinchipe: border, crest of Cordillera Oriental, 
2,840 m, 28 Jan 1984 (fl),J. Luteyn & E. Cotton 1 1295 (NY, QCA); W slopes of Cordillera 
del Condor and NW slopes of Nudo de Sabanillas, around Tambo Cachiyacu, ca. 2 km SE 
of Yangana, 2,000-3,000 m, 19 Oct 1943 (fl, fr),./. Steyermark 54800 (NY, U); S of El 
Playon de San Francisco, slopes of Cerro Mirador, 3,300-3,600 m, 29 Dec 1980 (fl, fr), L. 
Hotm-Nielsen et al. 29949 (AAU),J.Jarmtito et al. 3929 (AAU, QCA). PERU. Amazonas: 
Prov. Luyas, Dtto. Camporredondo, Ancxo Tullanaya, Cerro Wicsocunga, 06° 05' 35" S, 
78° 19' 56" W, 3,075 m, 7 Dec 1996 (pist. fl),/ & L. Campos 3121 (BRIT, MO, USM). 
Cajamarca: Jaen, SW of Querocotillo, 3,150 m, Aug 1915 (bisex. fl, fr), A. Weberbai/er 
7168 (F, G, GH). Cusco: La Convencion, 2,800 m, 9 Jul 1968 (bisex. fl, fr), T. Dudley 
10910 (NA), 10 Jul 1968 (bisex, fl, fr), T. Dttdley 10922 (E, NA), T. Dudley I0931B (F, 
NA). Huanuco: Prov. Huanuco, 45 km on rd. from Huanuco to Tingo Mari'a, trail on S 
sideofCarpish Tunnel, 09° 42' S, 76° 05' W, 2,400 m, 3 Mar 1985 (bisex. fl), C. Todzia & 
B. Stew 2740 (F, TEX, USM). 

As was stated previously (Pipoly 1987), Cybianthus marginatus is most 
closely related to C. liueat/is (Benth.) Pipoly, previously known only from 
the contiguous Guayana Floristic Province, because of its (synapomorphic) 
flexous inflorescences and biseriate ovules. However, Cybianthus niargtnatus 
IS easily distinguished from C. liutatiis by its glabrous, ridged, verrucose- 
papillate branchlets, oblong, elliptic or ovate leaves, obnapiform pistil and 

PiPOLY, Cybianthus in Ecuador and Peru 117 

ellipsoid fruits. The verrucose-papillate branchlets, leaf and fruit shape are 
unique (autapomorphic) features within the subgenus. 

Cybianthus margtnatns is the most variable species of the subgenus, con- 
taining one-third of the names attributed to Grammadmta as taxonomic synonyms. 
Variation in leaf size and punctation and quantitative floral variation have 
led to much overdescription. A full disctission of synonymy and variation 
was provided by Pipoly (1987). Cybianthus marginatus is most closely re- 
lated to C. limatus by virtue of its flexuous inflorescences and biseriate ovules. 
However, Cybianthus marginatus is easily distinguished from C. lineatus by 
the glabrous, ridged, verruose-papillate branchlets, oblong, elliptic or ovate 
leaves, obnapiform pistil and ellipsoid fruits. 

29. Cybianthus lineatus (Benth.) Pipoly (Fig. 7E), Mem. New York Bot. 
Gard. 43:64. 1987. Gnimmcuknhi limata Bentham, PI. Hartw. 218. 1846. Type: 
VENEZUELA. ["GUYANA"}, houvar: Savannas near Roraima, 1843 (bisex. fl, fr), R. 
Schonih/a-gk 647/992 (i lOLOTYPr,: K; isotypes: B, BM, G-DC, G-DEL, P, U, W-2 sheets). 

Terrestrial shrub to 1 .5 m tall, the trunk pachycaulous. Branchlets terete, 
(2.5-)3— 4(— 6) mm diam., densely red glandular-papillate apically. Leaves 
coriaceous, symmetrical, oblanceolate,(1.2-)1.6-3(-3. 9) cm long,(0.4-)0.6- 
1 (—1 .2) cm wide, apically acute, basally subauriculate, the margin flat. Inflorescence 
erect, flexuous, at times reduced to a simple dichasium, 0.6— 1.0(— 2.5) cm 
long, sparingly glandular-papillate; floral bracts widely ovate, 0.8—1 . 1 mm 
long, 1.1—1.4 mm wide, apically acute, the margin entire, glabrous. Flow- 
ers chartaceous, 5-6(-7)-merous; calyx 1.1-1.6 mm long, the tube ca. 0.1 
mm long, the lobes wide-triangular to deltate, (0.8-)l-1.5 mm long, (0.7-) 
1-1 .2(-l .4) mm wide, apically acute, conspicuously black punctate and punctate- 
lineate, the margins entire, glandular-ciliolate; corolla (1.8— )2— 2. 5(— 3) mm 
long, the lobes widely ovate, (1-)1 .3-1 .6(-2) mm long, (1-)1 .3-1.7(-2) 
mm wide, obtuse to emargmate, inconspicuously punctate medially, the 
margins irregular, entire; staminal tube carnose, conspicuous, (0.6— )0. 8—1 
mm long, the anthers sessile, alternate with prominent fleshy lobes, quad- 
rate, 0.4-0.6 mm long and wide, apically rounded, prominently black punctate 
dorsally; pistil ellipsoid, (0.9-)l-l .2(-l .4) mm long, (0.8-)l-l .3 mm diam., 
the ovary 0.8—1 mm long, glabrous to translucent glandular-lepidote apically, 
ovules 2-3, when more than 2, biseriate, the style (0.1-)0.2(-0.3) mm long, 
glabrous. Fruit ohowo'id, (3-)4-5 mm long, (2-)2.5-3 mm diam., purple, 
than black at maturity, the punctations green, prominent. 

Distribution. — Formerly thought to be endemic to the Guayana High- 
land, in open savannas on tepuf summits throughout Pantepuf (Mayr & Phelps 
1967), at 1,400-2,850 m elevation, but now known elsewhere only from 
the collection cited below. 

Ecology and conservation status. — Cybianthus lineatus grows in fully exposed, 

118 SiDA 18(1) 

dry montane "shrub savannas" (Huber 1995) in shallow sand over sand- 
stone throughout Pantepui. Its occurrence in Pajonal ("jalca") vegetation 
in Peru gives reason to expect it in other places, especially in the Cordillera 
del CcSndor along the Ecuadorean/Peruvian border. Unfortunately, recent 
civil Luirest has prohibited collection in that area. In the majority of the 
range lor the species, Cybianthus lineatus is not threatened. 

Etymology. — The specific epithet refers to the numerous and prominent 
punctate-lineations of the abaxial leaf surface. 

Specimen examined. PERU. Pasco: Oxapampa l-'rov., ( ^rro Pajonal, 29 km from Oxapampa, 
2,680 m, D. Smith & [-Mter 2'>09 (F, MO, USM). 

Cybianthus lineatiis is most closely related to C marginatum (Benth.) Pipoly, 
by virtue of its flexuous inflorescence and biseriate ovules (Pipoly 1987). 
However, Cyb'mnthin lineatus may be easily recognized by its pachycaulous 
trunk, densely red glandular-papillate branchlet apices, ellipsoid pistil and 
obovoid fruits. It is the only species in the subgenus with parenchyma in- 
stead of aerenchyma in the cortex, the only one with bifacial palisade layers 
in the leaf, and the only one with a pachycaulous trunk. None of these 
morphological peculiarities are unexpected given its drier, wind-swept habitat. 

30. Cybianthus magnus (Mez) Pipoly, Mem. New York Bot. Gard. 43:55. 1987. 

Facultative epiphytic shrub or tree to 7 m tall. Branchlets terete, smooth, 
glabrotis. Leaves chartaceotis, asymmetrical, narrowly oblanceolate, oblan- 
ceolate or narrowly obovate, (4.5-)5.2-15 cm long, (l.()-)2. 1—5.2 cm wide, 
apically acute to abruptly acuminate, tapering abruptly or gradually to base, 
bearing hydropotes above or not, conspicuously black punctate and punc- 
tate-lineate or inconspicuously pellucid to orange punctate below, the margins 
entire, flat, or subrevolute. Inflorescence lax, straight, (1.5-)2-8(-l 1.5) cm 
long, slender, densely glandular-granuiosc and papillate, prominently black 
punctate-lineate or conspicuously orange to brown ptinctate-lineate; floral 
bracts ovate, widely ovate or deltate, (0.7— )1. 1-2.2 mm long, (().6-)l .3-2 
mm wide, apically acute to acuminate, prominently black or orange punc- 
tate and punctate-lineate, the margins erose and glandular-ciliate; pedicels 
1.0— 2. 2(— 5.5) mm long in flower, the smaller ones accrescent to 4(— 6) mm 
long in fruit. Flowers chartaceotis, 5(— 6)-merous, whitish-green or pink to 
maroon; calyx lobes widely ovate to cielatate, (0.8— )1-1 .5(— 2) mm long, 
(0.8—) I -2.1 mm wide, apically acute to acuminate, prominently black punctate 
and punctate-lineate or orange to brown punctate, the margins erose to fimbriate 
and densely glandular-ciliate; corolla (1 .7-)2-2.6(-3) mm long, the stami- 
nate and bisexual maroon, the pistillate white, the lobes widely ovate, 1.1— 
1.6(-2.2) mm long, (0.9-)l . L-2.6 mm wide, apically obtuse to emargin- 
ate, rugose medially within, densely and prominently black punctate and 

PiPOLY, Cybianthus in Ecuador and Peru 119 

punctate-lineate or orange to brown punctate, the margins irregular, en- 
tire; staminal tube membranaceous, (0.4— )0. 7—1 (—1.2) mm long, lobate, 
the lobes 0.1-0.2 mm long, the anthers sessile, alternate with the lobes, 
ovate to quadrate, (0.3-)0.4-0.6 mm long, 0.4— 0.6(— 7) mm wide, apically 
rounded or emarginate, the connectives epunctate ventrally, prominently 
black or orange punctate dorsally; pistil obnapiform, 1-1.2 mm long, 0.9— 
1.5 mm diam., the ovary 0.6— 0.9(— 1 -3) mm long, densely translucent glandular- 
lepidote, the ovules 2-4(-5), uniseriate, the style 0.3-0.5(-0.7) mm long, 
glabrous. Fr//i/ obovoid, 2.5—3.5 mm long, 2-3 mm diam. when dried, pink, 
then purple-black or white, then lavender at maturity, prominently black 
punctate-lineate or orange punctate and punctate-lineate when dried. 

Distribution. — Cybianthus ma gnus occurs in the Andes of Venezuela southward 
to Peru as an epiphyte in the cloud forest zone below subparamo thickets 
and as a facultative epiphyte in montane and elfin "ceja" forests, from 1 ,100— 
3,500 m. It is also known from the Serranfa de Turumiquire, in the states 
of Monagas, Sucre and Anzoategui, Venezuela, at 2,000-2,400 m. 

Etymology. — The specific epithet refers both to the large, branchlets, often 
appearing succulent, as well as the large leaf size found in some populations. 

As shown by Pipoly (1987) Cybianthus magnus is most closely related to 
C. parasiticus (Sw.) Pipoly from the Lesser Antilles by its chartaceous co- 
rolla rugose medially within, asymmetrical leaves, and obovoid fruits, but 
is easily separated from it by its erose and fimbriate calyx lobes, lobate staminal 
tube and sessile anthers. Cybianthus magnus superficially resembles C. marginatus 
(Benth.) Pipoly, but may be easily separated by its subsucculent smooth 
stems, and obovoid fruits 

Both subspecies are known from Ecuador and Peru, with one region of 
apparent sympatry on the eastern slopes of the Cordillera Oriental in Ecua- 
dor and adjacent northern Peru. However, they appear to be separated by 
habitat. In Ecuador and Peru, subspecies magnus appears to be restricted to 
closed cloud forests, while subspecies asynmietricus is found in open mon- 
tane forest and elfin ("ceja") forest. The salient features of each are summa- 
rized below. 

30a. Cybianthus magnus (Mez) Pipoly subsp. magnus (Fig. 8E). Mem. 
New York Bot. Gard. 43:56. 1987. Gnjmvuidefiui magnaMez'm'Engi.^V'Aiinz.tnr. 
IV. 236(Heft 9):231. 1902. Tvph: COLOMBIA. Santander del Norte: Ocana to 
Pamplona, 2,000-2,500 m, 4 Mar 1S79 (tl), W. Kalhreyer 1087 (hou)TYFh: K). 

Grammadenia oxygyna Cuatrec, Revista Acad. Colomb. Ci. Exact 8:321. 1951. T'^te: 
COLOMBIA. Valle di;L Caica: Cordillera Occidental, W slope, bank of Rio Digua, 
left side, Piedra de Moler, 900-1 , 100 m, 20 Aug 1943 (fl, fr),/ Cuatrecasas 14947 (lcc;to- 
TYPE by Pipoly (1987): F, NY Neg 12136; lsole(;totypes:COL-3 sheets, F, U, US). 

Facultative epiphytic shr/^b or tree to 7 m tall, 7-15(-30) cm diam., the 

120 SiDA 18(1) 

canopy often bowl-shaped. Branchlets (4.0— )5. 0—7.0 mm diam. Leaves not 
bearing hydropotes above, conspicuously black punctate and punctate-lineate 
below. Inflorescence rachis prominently black punctate and punctate-lineate. 
Flowers with perianth whitish-green, prominently black punctate and punctate- 
lineate; anthers ovate, rounded apically, the connectives prominently black 
punctate dorsally. Fr//it purple-black at maturity, prominently black punc- 
tate-lineate when dried. 

Distribution. — Subspecies magniis occurs from the Serrania de Turumiquire 
(states of Anzoategui, Monagas, Sucre), and in the Andes, from Venezuela 
southward through Colombia to and Ecuador to Peru. 

Ecology and conservation status. — Subspecies magnus occurs as an epiphyte 
in closed cloud forests, especially those below subparamo thickets. It is a 
relatively rare, but widely distributed subspecies, and is increasingly en- 
dangered owing to habitat destruction. 

Etymology. — The epithet refers to the leaf size and stem succulence, a novelty 
within the subgenus. 

Specimens examined. ECUADOR. Loja: Between Nudo de Sabanillas and Rio Cachiyacu 
at Tambo Cachiyacu, 3,()()()-3,5()() m, 1 7 Oct 1943 (IDJ. Stuymuark 53584 (NY). Zamora- 
Chinchipe: Nangaritza Cant6n, ridge crest of Cordillera del Condor, above Pachiciitza, on 
disputed Peru-Ecuador border, ()4° 06' S, 78" 35' W, 1 ,800 m, 5 Dec 1990 (stam. (1), D. 
Neill & W. ?alcicm 9318 (MO, QCNE). PERU. Cajamarca: Prov. San Ignacio, path at the 
border of "La Union," 2,200 m, 1 Nov 1995 (stam, H), C. D/az & A. Torres 7805 (BRIT, 
MO, USM). 

30b. Cybianthus magnus (Mez) Pipoly subsp. asymmetricus (Mez) Pipoly 
(Fig. 8F), JVLem. New York Bot. Card. 43:57. 1987. Cr^immckuicicisyiuuietriai 
Mez, Bull. Herb. Boissier ser 2, 5:246. 1905. T^'Pi;: PERU. Loiurro: Cerro de Ponasa, 
1 ,300 m, Mar 1903 (fr), E. Ule 6792 (l.i^CTOTYPi.; by Pi]M)ly 1987: HBG; i.solf.ctotyprs: 
F, G, K, L). 

Gramiiiaden'ui niacrocarpci Lundell, Wrightia 5:292. 1976. Type: ECUADOR. Napo: 17 
km W of Lumbaque, 70-73 km W of Lago Agno, 1,130 m, 4 Nov 1974 (bisex. fl, 
fr), A. Gaitry 1241') (hoi.otypp.: LL-TRX; isotyph.s: MO, S). 

Facultative epiphytic .t/ir///; to 2 m tall, 7 m diam., the canopy open, conical. 
Brancblets 2.5— 3. 5(— 4.0) mm diam. Leaves bearing hydropotes above at least 
proximally, inconspicuously pellucid to orange punctate below. Inflorescence 
rachis conspicuously orange to brown punctate-lineate. flowers with calyx 
greenish-pink, the staminate and bisexial with corolla maroon, the pistil- 
late with corolla white and prominently orange punctate and punctate-lineate 
lobes; anthers quadrate, apically emarginate, the connectives prominently 
orange punctate dorsally. Fruit white, then lavender at maturity, prominently 
orange punctate and punctate-lineate when dried. 

Distribution. — Cybianthus magnus subsp. asymmetricus occurs as an obligate 
epiphyte, growing on detritis in open montane forests and elfin forests, from 
the Darien of Panama to Cusco, Peru, from 1 ,000-2,000(-2,700) m. 

PiPOLY, Cybianchus in Ecuador and Peru 121 

Ecology and conservation statns. — Subspecies asymmetricus grows as an obli- 
gate epiphyte, growing on rocks with deep organic detritis and large, moss- 
covered trees. With increasing levels of disturbance, it is being threatened. 

Etymology. — The epithet refers to the asymmetric shape of the leaf blades. 

Specimens examined. ECUADOR. Carchi: Canton Tulcan, Parroquia Tobar Donoso, 
Reserva Indigena Awa, Centre El Baboso, 00" 53' N, 78° 25' W, 1,800 m, 17-27 Aug 
1992 (fr), G. Tipaz etal. 1 741 (BRIT, MO, QCNE); Parroquia el Chical, Centro San Marcos, 
01° 06' N, 78° 14' W, 900-1,100 m, 20-30 Apr 1993 (A), P- Mendez et al. 341 (BRIT, 
MO); Cerro Golondrinas, valley bottom ca. 1.5 km NNE of summit, 00° 51' 52" N, 78° 
08' 10" W, 2,750 m, 25 Jul 1994 (stam. fl), B. Boyle etal. 3450 (BRIT, QCNE, MO). PERU. 
Amazonas: Prov. Bagua, Imaza, Nuevo Samaria (anexo de UVT), 18 Mar 1995 C. Diaz et 
cil. 7385 (BRIT, HUT, MO, USM), Cusco: La Convencion, Cordillera Vilcabamba, ca. 1/2 
way between Camps 2 1/2 and 3, 1,980 m, 1 Jul 1968 (bisex. fl, fr), T. Diulley 1()66S (F, 
NA), 1,800 m, 24 Jul 1968 (bisex. fl, fr), T. Diulley 1 1324 (NA). Huanuco: Prov. Pachitca, 
region of Pucallpa, W part of Sira Mountains and adjacent lowland, ca 24 km SE to 26 km 
ESE of Puerto Inca, next to Campamento Pato Rojo, 09° 27' S, 74° 46' W, and along crest 
after Campamento Peligrosol,600 m, 14 Apr 1 988 (fr), B. Wallmfer 18-144H8 (BRIT, MO, 
W, WU). Pasco: Oxapampa, Cordillera Yanachaga, Cerro Pajonal, chacos, 12 km SE of 
Oxapampa, 2,700-2,800 m, 7 Oct 1983 (bisex. fl), R. Foster 9013 (MO, NY, USM). 

Cybianthus magnus subsp. asymmetricus is notable for its stems with angu- 
lar coUenchyma in the pith, well-developed aerenchyma in the inner cor- 
tex, and tangential collenchyma in the outer cortex. It is separated from 
subspecies magnus by its inconspicuous or orange punctate leaves, orange 
punctate-lineate inflorescence rachis and perianth, quadrate, emarginate anthers 
and open montane and elfin forest habitat. 

Grammadenia macrocarpa Lundell is notable only for its large, bright or- 
ange fruits and smaller flowers. I examined one fruit from each of the isotypes 
of G. macrocarpa and found an insect larva in each one, accounting for the 
size and peculiar morphology. 

VIII. Cybianthus Mart, subgenus Cybianthus. Cyhianthus sect. Encylnanthus 
Miq. in Mart., Fl. Bras. 10:292. 1856. Cybiautbi/s sett. Cybumthoidei Miq. in Mart. 
Fl. Bras. 10:292. 1856. Typh SPECrES. Cyhianthns pendulifiorus M-An. 

Peckia Veil., Fl. Flum. 1:5 1. 1825., nom. rej. 

Terrestrial dioecious shrubs or small trees. Roots positively geotropic. Bark 
mostly smooth, gray or sometimes brown, cracking longitudinally on older 
parts. Trunks distinguishable, normally more or less terete, leptocaulous, 
growth dynamics corresponding to Rauh's Model (Halle et al . 1 978). Branchlets 
thin to moderately thick, terete, trigonal, or ridged, rufous stellate or den- 
droid tomentose or rarely, with rufous, subsessile covering lepidote scales. 
Cataphylls and pseudocataphylls absent. Leaves alternate, or pseudoverticillate, 
supervolute, exstipulate, petiolate; blades epunctate or variously black or 
red punctate, the punctations at times prominent. Inflorescence a simple, lat- 
eral (axillary) raceme; peduncle 1-5 mm long, the rachis straight, minutely 

122 Si DA 1,S(1) 

rufous glandular-papillate, glandular-granulose or lepidote, glabrescent or 
persistent; floral bracts solitary, at pedicel basally, ovate to lanceolate, per- 
sistent; pedicels cylindrical to clavate, or obconic, minutely glandular-papillate, 
glandular-granulose or lepidote, accrescent in fruit. Flowers unisextial or bisexual, 
4-merous; perianth lobes imbricate or rarely valvate, prominently, conspicuously 
or inconspicuously punctate and/or punctate-lineate; calyx cotyliform, the 
lobes erect, at times reflexed in fruit, the margins irregular or regular, en- 
tire or minutely crenulate, glandular-ciliate; corolla rotate, to subrotate, the 
tube short, glabrous or glandular-granulose, at times papillate, the lobes 
glabrous without, glandular-granulose and/or glandular-papillate throughout 
within, the margins regular or irregular, entire or minutely crenulate, gla- 
brous, glandular-granulose or papillate along the margins; stamens and 
stami nodes similar, developmentally adnate to the corolla tube to form an 
inconspicuous tube, the stamens thus appearing epipetalous, apical free portion 
of the filaments present or absent, when present up to 3 times longer than 
the anthers, the anthers basifixed, appearing sessile or on minute apically 
free filaments, quadrate or ovate, apically emarginate, rounded, acute, api- 
culate or truncate, basally truncate or subcordate, dehiscent by apical pores, 
the pores at times confluent, the connective epunctate or conspicuously or 
prominently punctate, at times glandular-papillate; pistil and pistillode similar, 
conic, ellipsoid, or rarely, obturbinate, the ovary terete, lobed or costate, 
sparsely to densely translucent-lepidote, the placenta subglobose, (l-)2- 
3(— 4) ovulate, the ovules half-immersed in the basal placenta, uniseriate or 
biseriate, the style long-attenuate, the stigma punctiform, lobed or capi- 
tate-lobed. Fr//// drupaceous, depressed globose, the exocarp sometimes fleshy, 
the mesocarp and endocarp stony, prominently punctate-lineate, white, red 
or purple-black at maturity, one seeded, the testa corrugate, the embryo 
cylindrical, transverse. 

A SLibgenLis of 50 species in tropical South America, with the largest 
concentration ol species in the Amazon Basin and adjacent Guianas, Planalto 
and coastal Brazil. Subgenus Cybianthia is by far the most complicated taxonomic 
group within the genus and is known from rather incomplete material. Collection 
of more material is hampered by the fact that the populations studied here- 
tofore have revealed population densities lower than any other subgenus. 
So far, 13 species are known from Ecuador and Peru, but with additional 
collections, we may expect to find Cybianthus lanceolatm Pax, and/or Cybtcinthns 
psychotriijoliiis (Rusby) Mez, both from nearby Bolivia, in southern Peru. 


1. Branclilecs subteretL- to ant^'iilate. 

2. Indorcscence spicarc- or SLibspicatc; calyx lobes inconspicuously or promi- 
nently orange piincrate; petioles pulvinace basally. 

PiPOi.Y, Cybianthus in Ecuador and Peru 123 

3 . Leaf blades inconspicuously pellucid punccate below; calyx lobes chartaceous 
or carnose, rounded or acute apically. 

4. Branchlets 4—6 mm diam.; leaf blades coriaceous, nitid above; peti- 
oles 2.2-3 cm loni;; inflorescence spicate, the pedicels obsolete to 
0.4 mm long; calyx lobes chartaceous, deltate, the margin entire, 

undulate, glabrous 31. C. incognitus 

4. Branchlet 2—3.9 mm diam.; leaf blades membranaceous, dull above; 
petioles 1.7-2 cm long; inflorescence stibspicate, the pedicels 0.6- 
0.9 mm long; calyx lobes carnose, ovate, the margin crenulate, glandular- 

ciliace 32. C. minutiflorus 

3. Leaf blades perpuncticulose below; calyx lobes membranaceous, 

subacuminate apically 33- C. huampamiensis 

2. Inflorescence racemose; calyx lobes densely and prominently black punc- 
tate or epunccate; petioles tapered, not pulvinate basally. 
5. Leaves alternate; calyx lobes acute apically, the margins hyaline; an- 
ther apically rounded, obtuse or acute apically, the pores not confluent. 
6. Pedicels 1.9-2.2 mm long; flowers erect; calyx carnose, 0.8-1 .1 mm 
long, the lobes ovate, abruptly constricted basally, densely and promi- 
nently black punctate, the margin irregularly serrate, punctate-lineate, 

glabrous 34. C. granulosus 

6. Pedicels 2.5-3.5 mm long; flowers nodding; calyx membranaceous, 
1.5-1.9 mm long, the lobes deltate to triangular, not constricted 
basally, epunctate, the margin entire, epunctate, minutely glandu- 

lar-ciliolate 35. C. flavovirens 

5 . Leaves pseudoverticiUate; calyx lobes rounded apically, the margins opaque; 

anthers truncate apically, the pores confluent 36. C. venezuelanus 

1. Branchlets terete. 

7. Branchlets thick, (6-)7-10 mm in diameter. 

8. Leaf blades subacuminate apically; calyx membranaceous or chartaceous, 
the lobes acute or acuminate. 

9. Leaf blades nitid and perpuncticulose above, 12—13.5 cm wide, the 
secondary veins prominently raised above and below, basally trun- 
cate, auriculate; petioles 0.5-1.4 cm long; pedicels 0.3-0.5 mm long. 

37. C. grandezii 

9. Leaf blades sordid and epunctate above, 2.1—5 cm wide, the second- 
ary veins deeply impressed above, prominently raised below, basally 
cuneate; petioles 2.1-5 cm long; pedicels 0.8-1.4 mm long. 

38. C. jensonii 

8. Leaf blades abruptly acuminate apically; calyx coriaceous, the lobes 

rounded 39- C. fosteri 

7. Branchlets thin, 1.5-3.5 mm in diameter. 

1 0. Leaf blades apically subacuminate to acuminate; calyx lobes acuminate 
or attentuate, the margin erose, short glandular-ciliate. 
1 1 . Leaf blades chartaceous to coriaceous, somewhat to very nitid 
above and below, the midrib raised above, decurrenr to base 
of petiole; petioles short-pulvinate basally. 
12. Leaf blades elliptic, (4-)5.5-7(-10.8) cm wide, apically 
long-acuminate, basally cuneate, the tertiary veins prominently 
raised, inconspicuously pellucid-punctate below; petioles 
canaliculate; pedicels cylindrical in fruit 40. C. resinosus 


SiDA 18(1) 


FKi. 22. Cyhhiiitbiis ntoiv^iiitiis I'lpoly. A. Habit, showing rrit^onal branchler. B. Abaxial leaf 
surface, showing miniiCL- scales. C. Portion of infrLictcscencc, showing delcate calyx lobes 
with entire margins. D. Branchler apex, showing dendrojd and stellate tomentum. E. Por- 
tion of staminate spike, showing conspicuous staminal tube, and obcordate anthers with 
SLibapical non-confluent pores. A, B, O, drawn from Gt:)it)-y et al. 2291 1. C, drawn from 
B(/r/x)//r 2567 . E, drawn from holotype, by Lintia EUis. 

Pipon', Cybianthus in Ecuador and Peru 125 

12. Leaf blades very narrowly oblanceolate or oblong, 2— 4(— 5) cm 
wide, apically and basally long-attenuate, the tertiary veins 
inconspicuous, conspicuously black or red punctate and 
punctate lineate below; petioles marginate; pedicels obconic 

in frti i t 4 I.e. fuscus 

1 1. Leaf blades chartaceous, di.ill green above and below, the mid- 
rib impressed above, not decurrent on the petiole; petioles gradually 

tapering to base, withotit ptilvinus 42. C. cyclopetalus 

10. Leaf blades apically acute; calyx lobes obtuse, the margin crenu- 

late, long ciliate 43. C. penduliflorus 

31. Cybianthus incognitus Pipoly, sp. nov. (Fig. 22). Typi;: PERU. Amazonas: 
Rio Santiago Valley, 03° 50' S, 77° 40' W, QuebradaCaterpiza, 2-3 km from Caterpiza 
settlement, primary foresr, 200 m, tree 9 m tall, 12 Dec 1979 (stam. fl), S. Tiinqui 
289 (holotype: MO; isotypes: USM, NY). 

Ob folia coriacea oblanceolata desuper nitida subter pallida anthera filamenta 3-plo breviores 
C. prieuro valde arete affinis sed ab ea petiolis canaliculatis (nee marginatis), ramulis adpresse 
dendroideo- et stellate- (nee erecte dendroideo-) tomentellis, pedicellis 0—0.4 (non 0.8— 
1.4) mm longis, lobiscalycinis inconspicuepellucido- (non manitesre aero-) punccatis, lobisw 
coroUinis ovatis (nee stiborbicularis) pistillodio globoso (nee conico) denique fructu laevi 
statim recognitur. 

Tree to 9 m tall, at times flowering precociously {P. Barbour 2405). Branchlets 
subterete to trigonal, 4—6 mm diam., appressed rufous dendroid and stel- 
late tomentose. Leaves pseudoverticillate; blades coriaceous, oblanceolate, 
(16— )20— 25(-31) cm long, (5— )6.5— 8.5(— 10) cm wide, apically acuminate, 
basally cuneate, decurrent on the petiole, nitid above, pallid below, the midrib 
slightly impressed above, prominently raised below, the secondary veins 9— 
12(— 15) pairs, slightly raised above, prominently raised below, nitid above, 
pallid and minutely rubiginous lepidote below, the pellucid punctations 
inconspicuous, the margin entire, irregular, flat; petioles canaliculate 2.2— 
3 cm long, pulvinate, sparsely pubescent at first, glabrescent. Staminate inflorescence: 
an erect, dense spike, (6— )8— 9-5 cm long, the rachis green, sparsely den- 
droid pubescent, glabrescent; floral bracts lanceolate, 0.5-1 mm long, apically 
attenuate, sparsely pubescent, early caducous; pedicels obsolete to stoutly 
cylindrical, 0—0.4 mm long, glabrate. Staminate flowers 4-merous, yellow, 
chartaceous; calyx cotyliform, 0.6—0.9 nxm long, the tube 0.2 mm long, 
the lobes deltate, 0.4—0.7 mm long and wide, apically acute, inconspicu- 
ously pellucid punctate, the margins scarious, entire, epunctate, undulate, 
glabrous; corolla subrotate, 1.4—1.6 mm long, the tube 0.2—0.3 mm long, 
the lobes ovate, 1.2—1.4 mm long, 0.4—0.6 mm wide, apically acute, gla- 
brous without, densely glandular-granulose within, inconspicuously pel- 
lucid punctate, the margin opaque, densely glandular-granulose, entire; stamens 
1—1.2 mm long, the tube ca. 0.2 mm long, the filaments terete, 0.5—0.6 
mm long, slightly reflexed proximally, the anthers obcordate, 0.3—0.4 mm 
long, apically apiculate, basally cordate, dehiscent by small, subapical, ovate. 

126 SiDA IS(I) 

non-conHuent pores, the pores extendin^^ less than 1/2 anther length, the 
connective epunctate, densely and minutely rubiginous glandular-granulose 
dorsally; pistillode broadly subglobose, ca. 0.2 mm long, 0.4 mm diam., 
densely yellow glandular-papillate. P istillate inflorescence as in staminate but 
9—12 cm long; floral bracts lanceloate, 0.2—0.5 mm long, glabrate; pedicels 
obsolete to 0.4 mm long, glabrate. P isti I Ic/te flowers as in staminate, but ca- 
lyx 0.6—0.9 mm long, the tube 0.2 mm long, the lobes 0.4—0.7 mm long 
and wide, the margins opaque, entire; corolla, staminodes and pistil un- 
known. Fr///'/ dark purple at maturity, globose, 0.4—0.5 mm long, 0.4—0.6 
mm diam., smooth, incospicuously pellucid punctate. 

Distribution. — Upper Rio Santiago Valley and adjacent Serranfa de Bagua, 
Amazonas, 200—2,000 m, and Maynas Province, Loreto, Peru, along the 
Rfo Napo, at 120 m elevation. 

Ecology and conservation status. — Cyhianthi/s incognitas is mostly a ridgetop 
species in the cloud forests of Amazonas Department, occurring infrequently 
near the forest margin. The Upper Rio Santiago Valley and adjacent Serranfa 
de Bagua are known for their endemic species (Pipoly 1992b). However, 
one surprising collection was noted at 120 m elevation, from Caserio de 
Urcumiraho, in Maynas Province of Loreto Department, where vegetation 
normally associated with much higher altitudes occurs on the tops of un- 
dulating hills. Much more fieldwork will be required to better understand 
forest dynamics at the western limits of the Peruvian Amazon with the foothills 
of the Andes. 

Etymology. — The specific epithet refers to the fact that the plant was 
misidentified even to family for nearly twenty years, and was finally identified 
only when a flowering specimen was matched with the other fruiting specimens. 
The densely spicate infructescences with numerous fruits were heretofore 
unknown in the genus. 

Paratypi;s. PERU. Amazonas: Prov. Bagua, 12 km E of La Peca, cloud forest, 1700 m, 
20 Jun 1978 (fr), P. Barbour 2405 (AMAZ, F, MO, USM), 29 Jun 1978 (fr), P. Barhotir 
2367 (AMAZ, BRIT, F, MO, NY, US); Ca. 12-18 km E of La Peca m Serranfa de Bagua, 
cloud forest, 1 ,800-1 ,950 m, 14 Jun 1978 (fr), A. Gentry et al. 22839 (F, MO, USM), A. 
Gentry et al. 2291 1 (F, MO, USM). Loreto: Rio Napo near entrance to Isla Inayuga, 20 Sep 
1972 (fr), T. Croat 20328 (AMAZ, MO, USM); Caserio de Urcumirafio, Rfo Napo, 2 hours 
along trail from village to forest, 120 m, 8 Oct 1979 (stam. fl), C. Diaz & N. Jara>nillo 
1474 (AMAZ, BRIT, MO, USM). 

Cybianthus incognitus is appears to be most closely related to Cybianthus 
prieurii A. DC. of the Guianas, Venezuela and Brazil, because of the oblan- 
ceolate, highly nitid coriaceous leaf blades and the filaments three times 
longer than the anthers. However, Cybianthus incognitus is separated from 
C prieurii by its canaliculate petioles, dendroid and stellate tomentose branchlets, 
sessile to subsessile flowers (spicate inflorescences), inconspicuously pellu- 

PiPOLY, Cybianrhus in Ecuador and Peru 127 

cid-punctate calyx lobes, ovate corolla lobes, globose pistillode and smooth 
fruits. Cybianthus incognitus is unique within the subgenus by virtue of its 
densely spicate inflorescences, dendroid and stellate tomentum of the branchlets 
and inconspicuosly punctate calyx lobes. 

32. Cybianthus minutiflorus Mez, Repert. Spec. Nov. Regni Veg. 3:102. 
1 906. Type: PERU. Loreto: near Rioja, W of Moyobamba, 800-900 m, 8 Sep 1904 
(pist. fl, fr), A. Weberbauer 4699 (iiolotype: B -destr.; fragment, F; lectotype, here 
designated: F). Because the fragment at F contains floral and leaf material, and leave 
no doubt as to the identity of the species, in the absence of other duplicates, it is 
most appropriate to select this "clastotype" (a fragment taken with permission) as 
the lectotype. 

Tree to 3 m tall. Branchlets angulate, 4.5—6 mm diam., densely rufous 
stellate-tomentose, glabrescent. Leaves alternate; blades membranaceous, widely 
(rarely narrowly) oblanceolate, (l4-)19-28(-40) cm long, (4-^)9-1 2(-l 5) 
cm wide, apically acuminate, the acumen 1—1.5 cm long, basally cuneate, 
midrib slightly raised above, prominently raised and densely rufous tomentulose 
below, the secondary veins (9-) 12— 21 pairs, slightly raised above, promi- 
nently raised and sparsely rufous tomentulose below, smooth and glabrous 
above at maturity, sparsely rufous puberulent below, conspicuously pellu- 
cid punctate, the margin entire, flat; petioles canaliculate, 1.7—2 cm long, 
somewhat pulvinate, glabrous above, rufous tomentulose below, glabres- 
cent. Staminate inflorescence a simple, erect raceme, 4.5—9 cm long, the ra- 
chis densely rufous stellate-tomentose; floral bracts linear-lanceolate, 1 — 1.2 
mm long, 0.2—0.3 mm wide, apically attenuate, densely tomentose above 
and below, the margin entire; pedicels cylindrical, 0.6—0.9 mm long, densely 
tomentose, persistent. Staminate flowers pale yellow; calyx carnose, cotyliform, 
0.6—0.8 mm long, the tube ca. 0.1 mm long, the lobes widely triangular, 
0.5—0.7 mm long, 0.7—0.9 rnm wide, apically acute, densely and promi- 
nently orange punctate, sparsely rufous puberulent, the margin irregular, 
opaque, somewhat crenulate, minutely glandular-ciliolate; corolla carnose, 
subrotate, 1.2—1.4 mm long, the tube 0.3—0.4 mm long, the lobes widely 
triangular, 0.9—1.1 mm long, 1.1 — 1.2 mm wide, apically obtuse, densely 
and prominently orange punctate medially without, densely glandular-granulose 
medially and above anther within, the margin opaque, glandular-granulose, 
entire; stamens apparently sessile at corolla tube apically, the anthers widely 
obcordate, 0.4—0.5 mm long, 0.7—0.9 rnm wide, apically acute, basally cordate, 
the thecae moderately yellow glandular-granulose, the connective promi- 
nently red punctate; pistillode conic, 0.3—0.4 mm long, 0.1-0.2 mm diam., 
the stigma glandular-papillate. Pistillate inflorescence as m staminate but (4—) 
8—13 cm long; floral bracts 0.6—1 mm long, 0.1-0.2 mm wide, apically 
attenuate, densely tomentose above and below, the margin entire; pedicels 
0.6—0.8 mm long, accrescent in fruit to 1.8 mm long. Pistillate flowers as in 

128 SiDA 18(1) 

staminate but green; calyx 0.8-1 mm long, the tube ca. 0.1 mm long, the 
lobes 0.7-0.9 mm long, 0.9-1 mm wide; corolla 1-1.2 mm long, the tube 
0.2-0.3 mm long, the lobes 0.8-1.0 mm long, 1-1.1 mm wide; staminodes 
as in stamens but antherodes obcordate, 0.2-0.3 mm long, 0.2—0.3 mm 
wide; pistil cylindrical, 0.6-1 mm long, 0.2—0.3 mm diam., the ovary an- 
gular, 0.3-0.4 mm long, the style 0.3-0.6 mm long, the stigma subcapitate, 
4-lobed, the placenta cotyliform, ovules 2, naked. Fr//// depressed-globose, 
2.5-3.5 mm long, 4.5-5.5 mm diam., inconspicuously pellucid punctate, 
the exocarp thin. Bisexual inflorescence in staminate but 2-3(-5) cm long, 
the rachis moderately tomentose; floral bracts, 0.6-0.9 mm long, 0.1-0.2 
mm wide; pedicels 0.6-0.7 mm long. Bisexual flowers green; calyx 0.7-8 
mm long, the tube ca. 0.1 mm long, the lobes 0.6-0.7 mm long and wide; 
corolla 1.3-1.4 mm long, the tube 0.2-0.3 mm long, the lobes 1.1-1.2 
mm long, 1-1.1 mm wide; stamens identical to pistillate staminodes, but 
thecae full of pollen; pistil almost indistinguishable from pistillate flower 
except the stigma subcapitate, 3-4-lobed, ovule 1, naked. Fruit {horn bi- 
sexual flower) unknown. 

Distribution. — Endemic to the eastern slopes of the Andes and adjacent 
Amazonian Hylaea of Peru and adjacent Bolivia, 100-1200 m. 

Ecology and conservation status. — Cybianthus minutiflorus occurs in primary 
tall wet forest and premontane forest, on well drained white sands, known 
as varillal in Peru. These pockets of sandstone often alternate with rolling 
lateritic hills in the foothills of the eastern Andean slopes and adjacent Amazonia. 
The lowland forests where Cybianthus minutiflorus is occurs are also notable 
for their numerous lianas. Cybianthus minutiflorus is a rare species and should 
be considered threatened. 

Etymology. — The specific epithet refers to the extremely small flowers, 
some of the smallest in the subgenus. 

Local names and uses. — Peru: "taku kaspi" (Maynajivaro); leaves are boiled 
in water and the decoction drunk to treat stomach ache. 

Specimens examined. PERU. Amazonas: Prov. Bagua, Dtto. Imaza, Cerros de Putuim, 
05° 03' 20" S. 78° 20' 23" W, 350 m, 1 5 Jun 1996 (scam, fl), R. Vasquez tt al. 21187 (AMAZ, 
MO). Prov. Condorcanqui, Dtto. El Cenepa, NE region of Maranon Drainage Basin, Ri'o 
Cencpa, Comunidad Tutmo, 04° 33' S, 78° 10' W, 750 m, 22 Nov 1993 (fr), R. Vaujuezet 
al. 18520 (BRIT, HUT, JMO, USM). Cusco: Quipicanchi Prov., Camanti, Manirf, along 
Rio Maniri and along the trail to Quebrada Garrote, 13° 71' S, 70° 45' W, 720 m, 8 Sep 
1990 (bud), At, 71)/iatia922 (CUZ, MO, USM). Huanuco: Rio LluUa Pichis watershed, 
Cerros del Sira, 1,290 m, 17 Jul 1969 (fr),/ Wolfe 12346 (F, NA, US). Loreto: Prov. Loreto, 
Pampa Hermosa and vicinity, Rio Corrientes, 1 km S of junction with Rfo Macusari, 03° 
15' S, 75° 50' W, 160 m, 3-20 Dec 19S5 (fr), W. Lewis et al. 10312 (BRIT, MO); Prov. 
Maynas, Dtto. Las Amazonas, Explornapo Camp, near Sucusari, along Rio Napo, 03° 20' 
S, 72° 55' W, 100-140 m, 3 Mar 1991 (ster.),J, Pipoly et al. 14174 (MO, UNAP); Dtto. 
Iquitos, Allpahtiayo (IIAP), Permanent inventory, 04° 10' S, 73° 30' W, 150 m, 25 Mar 
1992 (ster.), R. Vasquez et al. IS 1 63 (BRIT, MO, UNAP). Pasco: Oxapampa, Rd. in con- 
struction between Oxapampa and Villa Rica, km 7, 10° 37' S, 75° 20' W, 2,100 m, 4 Jan 

PiPOLY, Cybianthus in Ecuador and Peru 



Fig. 23- Cybianthus huampamiensis Pipoly. A. Habit, showing pulvinate petioles. B. Abaxial, 
densely perpuncticulose leaf surface. C. Portion of staminate inflorescence, showing subspicate 
habit, and calyx erose-dentate and glandular-ciliate. D. Staminate flower, showing abruptly 
constricted corolla lobe base. E. Fruit and calyx. F. Branchlet apex. G, Pistillate flower, 
showing constricted corolla lobe base. A— B, drawn from Kayap 783 ■ C~D, drawn from Kayap 
982. E-G, drawn from Kayap 933- Figure drawn by Linda Ellis. 

130 Sum 18(1) 

1984 (bisex. fl), R. Foster et al. 787 7 (F, MO, USM); Oxapampa, Pichi's Valley, San Marias 
Ridge, 1 0—1 2 km SW of Puerto Bermudez, above Sta. Rosa de Chivis, trail to Loma Linda, 
10° 20' S, 75° 00' W, 1,000 m, 29 Sep 1982 (fr), R. Foster 8624 (R MO, USM), (stam. tl), 
R. Foster H981 (BRIT, MO, F, NY, US, USM). San Martin: Chazuta, Rio Huallaga, 260 m, 
Mar 1935 (stam. fl), G. King 3981 (F, MO, S, US); Prov. Mariscal Caceres, Dtco. Tocache 
Nuevo, Isla de Pucunchu, right bank of Rfo Huallaga, 3 Apr 1971 (lr),_/. Schunke 4779 (F, 
MO, NY, US); W of bridge, 700-800 m, 16 Dec 1971 (fr),/ Schunke' 37 37 (F, NY, MO, 
US). Ucayali: Prov. Coronel Portillo, Plantacion Margarita, near Loreto border, 1,500- 
1,600 m, 14 Aug 1946 (fr), R. Ferreyra 1040 (US, USM). BOLIVIA. Pando: Prov. Madre 
de Dios, Mobil Oil Site, 12° 10' S, 67° 15' W, 170 m, 20-25 Aug 1992 (fr), T. K/I/een 
4449 (BRIT, LPS, MO). 

Cyhianthus minutiflorus is most closely related to C. granulosus Pipoly by 
its densely rufous furfuraceous-lepidote branchlets, erect, carnose, densely 
and prominently black punctate perianth, and sessile anthers. However, the 
flat petioles, subacuminate leaf apices, short pedicels, opaque, crenulate and 
epunctate margins of the calyx lobes, and emarginate anthers easily distin- 
guish Cyhianthus minutiflorus. 

33. Cyhianthus huampamiensis Pipoly, sp. nov. (Fig. 23). Type: PERU. Ama7.ona.s: 

Quebrada chigkan entsa, Rfo Cenepa, 300 m, 9 Jun 1973 (stam. fl), F. Ancuash 388 
(holotype: MO; isotypps: NY, USM). 

Propter folia elliptica lanceolata vel oblanceolata, longipetiolata equilaterale vel inequilarerale 
secus margines irregulares, rhachides inflorescentiares graciles, flores erectes deminutosque, 
necnon Irutos minores, ad as]->ecru primo intuito C. resinoso arete similans sed ab ea laminis 
membranaceis (non renuiter coriaceis), utrinque sordidis (nee nitidis) subter manii:este 
prominenteque atro-perpuncticulosis (nee epunctatis), petiolis (1.5— )2— 2,5 (non 0.5—1 .4) 
cm longis, lobis calycinis translticentibus (non opacis) acuminatis (nee rotundatis), lobis 
coroliinis extus glandulari-granulosis (non glabris) acutis vel rotundatis (nee obtusis vel 
emarginatis) denique fructibus luteis (non atris) permanifeste distinguitur. 

Tree to 3(— 6) m tall. Branchlets subterete to angulate, (3-5— )4— 5 mm diam., 
densely rufous tomentose, glabrescent. Leaves psetidoverticiUate; blades 
membranaceous, elliptic lanceolate or rarely, oblanceolate, (12— )I6— 25(— 
31) cm long, (5.2— )7— 9(— 12)cm wide, apically long-acuminate, basally acute, 
not decurrent on the petiole, dull green above, pallid below, midrib im- 
pressed above, prominently raised below, the secondary veins 12—18 pairs, 
prominently raised below, glabrous above rufous puberulent below along 
the veins, prominently and densely perpuncticulose below, the margin en- 
tire, irregular, flat; petiole slightly canaliculate distally or flat, 2—2.5 mm 
long, thick and pulvinate, densely rufous puberulent at first, glabrescent. 
Stamina te inflorescence a lax, simple, subspicate raceme, (5—) 10— (18) cm long, 
sparsely rufous puberulent; floral bracts linear-lanceolate, 1—1.3 mm long, 
apically attenuate, densely rufous publerulent abaxially, the margin erose, 
persistent; pedicels cylindrical 0.8— 0.9 nim long, densely rufous pubescent, 
glabrescent. Staminate flowers yellowish to orange; calyx membranaceous, 
cotyliform, translucent, 1 .2—1.4 mm long, the tube ca. 0.2 mm long, the 

PiPOLY, Cybianthus in Ecuador and Peru 131 

lobes ovate, 1—1.2 mm long, 0.6—0.8 mm wide, apically subacuminate, densely 
and prominently orange punctate, glabrous, the margin hyaline, prominently 
erose-dentate, sparsely glandular-ciliate; corolla chartaceous, subrotate, 2.0— 
2.4 mm long, the tube 0.4—0.5 mm long, the lobes ovate, 1.6—1.9 mm long, 
1.2—1.4 mm wide, apically acute to rounded, sparsely glandular-granulose 
without and densely so throughout within, densely and prominently or- 
ange punctate, flat, the margin scarious, erose-denticulate and glandular- 
granulose; anthers apparently sessile at junction of corolla tube and lobe, 
very widely ovate, 0.4—0.5 mm long, 0.6—0.8 mm wide, apically obtuse to 
rounded, dehiscent by apical confluent pores extending ca. 2/3 length of 
anther, the connective epunctate, densely rubiginous glandular-papillate dorsally; 
pistillode conic, ca. 0.2—0.3 mm long and diam., hollow, glandular-papil- 
late. Pistillate inflorescence 3.?. in staminate but (5— )7— 9(— 14) cm long, densely 
rufous puberulent at first, glabrescent; pedicel 0.4—0.5 mm long. Pistillate 
flowers as in staminate but yellowish to orange; calyx 1—1.2 mm long, the 
tube ca. 0.2 mm long, the lobes 0.8-1 mm long, 0.4-0.5 mm wide; corolla 
1.8-2.2 mm long, the tube 0.3-0.4 mm long, the lobes 1.5-1.7 mm long, 
1.1-1.4 mm wide; staminodes as in stamens but antherodes 0.3—0.4 mm 
long, 0.5—0.6 mm wide, apically rounded to acute; pistil conic, ca. 1.3 mm 
long, the ovary 0.5—0.6 mm long 0.6-0.8 mm diam., translucent glandu- 
lar lepidote, the style short, to 0.2 mm long, the stigma 4-lobate, the lobes 
distally curved, glandular-papillate. Fr///V yellow, subglobose, 4—6 mm long, 
5—8 mm diam. inconspicuously pellucid punctate. 

Distribution. — Endemic to the Rfo Maranon, Rio Cenepa and Rfo Santiago 
drainage basins in the northwest corner of the Department of Amazonas, 
Peru, 200-550(-l,850)m. 

Ecology and conservation status. — Cybianthus huampamiensis occurs in the foothills 
of the premontane and lowland wet forest of the most underexplored area 
of the Peruvian Hylaea/Andean interface. Given that the region is a border 
area, and therefore, a priority for development, this species should be con- 

Etymology. — The epithet describes the place where the many of the col- 
lections were made, the Haumpami area of the Rio Cenepa Drainage Basin. 

Paratyprs. PERU. Amazonas: Prov. Bagua; ca. 12-18 trail km E of La Peca, Serranfa 
de Bagua, 1,800-1,950 m, 14 Jun 1978 (fr), A. Gentry et at. 22859 (F, MO, USM); Ri'o 
Cenepa, Quebrada tujushik entsa, 330 m, 18 Apr 1973 (fr), E. Ana/ash 274 (MO, USM); 
along Rio Cenepa, 350 m, 3 May 1973 (fr), H. Amuash 303 (MO, USM); Rfo Cenepa, Quebrada 
Idayua entsa, 400 m, 16 May 1 973 (fr), /;. Amuash 392 (BRIT, F, LL-TEX, MO, US, USM); 
Rfo Cenepa, Quebrada Wampusik entsa, "3 Aug 1974 (stam. fl), £. Ami/ash 731 (MO-2 
sheets, USM); Rfo Cenepa, Quebrada Cikan Inci, 250-330 m, 1 Jan 1973 (stam. fl), B. 
Bertni 779 (MO, VSM); 10 km N of Quebrada Huampami, 200-250 m, 24 Jul 1974 (stam. 
fl), B. Berlin 1760 (MO, USM); Quebrada Chigkui Shmuki Cenepa, 250 m, 1 1 Apr 1973 
(fr), R. Kayap 618 (MO, USM), 23 May 1973 (stam. fl), R. Kayap 783 (MO, NY, USM); 


SiDA l<S(l) 

Fk;. 24. Cyhui}itl)iis {i^ViDndoiiis Fipoly. A. Habic. B. Branchlet apex. (]. Abaxial leaf SLirhxce, 
showint; miiuicely riitous Icpitlocc indLimentum. D. Staminate flower. E. Staminate corolla. 
F. Sraminace tal\x. A-f], drawn from holotypie. D— F, drawn from \' Hinishikiit 1221 . 
Figure drawn by Linda Fllis. 

PiPOLY, Cybianthus in Ecuador and Peru 133 

Quebrada Etseketai, Rfo Cenepa, 250 m, 31 May 1973 (fr), R. Kayap 856 (MO, USM); 
Quebrada Wampushik entsa, 330 m, 13 Jun 1973 (pist. fl, fr), R. Kayap 933 (F, LL-TEX, 
MO, NY, USM); Huampami, Rio Cenepa, 200 m, 15 Jun 1973 (stam. fl), R. Kayap 982 
(LL-TEX, MO, NY, USM); Rio Cenepa, vicinity Huampami, ca. 5 km E of Chavez Valdivia, 
04° 30' S, 78° 30' W, 200-250 m, 31 Jul. 1978 (fr), A. Kujikat 30 (LL-TEX, MO, NY, 
USM), 12 Aug. 1978 (fr), A. Kujikat 291 (MO, NY, USM), 15 Aug. 1978 (fr), A. Kujikat 
395 (MO, USM); Mouth of Rio Santiago, without date (stam. fl), G. Tessmann 4439 (NY); 
Rio Santiago Valley, 03° 50' S, 77° 40' W, Quebrada Catetpiza, 2-3 km from Caterpiza, 
200 m, 28 Dec 1979 (stam. fl), S. Tunqui 488 (MO, NY, USM); Dtto. Imaza, Quebrada 
Kuzu, 1 hour walk from Comunidad de Cunchim, 370 m, 21 Jul 1994 (fr), C. Diaz et al. 
6930 (BRIT, HUT, MO, USM); Rfo Marafion Drainage Basin, Comunidad de Yamayakat, 
Rio Maranon, 04° 55' S, 78° 19' W, 600 m, 28 Jan 1995 (fr), £. Rodriguez 283 (BRIT, 
HUT, MO, USM), Quebrada Kusu-Chapi, 04° 55' S, 78° 19' W, 550 m, Feb 1995 (stam. 
fl), R. Vdsquez et al. 20045 (BRIT, HUT, MO), 320 m, 1 1 Mar 1996 (stam. fl), N.Jarami/h 
etal. 1351 (BRIT, HUT, MO). 

Cybianthus huampamiemis may at once be distinguished from all other species 
of the subgenus by its translucent calyx, externally glandular-granulose corolla 
and yellow fruit. This species is one of many endemic taxa known from this 
most underexplored area at the junction of the eastern Andean slopes with 
the Amazon Basin in northern Peru. 

34. Cybianthus granulosus Pipoly, sp. nov. (Fig. 24). Type: PERU. Amazonas: 
Rio Santiago Valley, 03° 50' S, 77° 40' W, Quebrada Caterpiza, 2-3 km from Caterpiza 
settlement, primary forest, 200 m, treelet 2 m tall, fls. brownish-green, 28 Nov 1979 
(stam. fl), S. Tunqui 161 (hoi.otype: MO; isotype: NY). 

Quoad folia alterna chartacea ad apices acuminata ad bases cuneataque, ramulos dense 
rufo-furfuraceo-lepidotos, flores erectos, carnosos, dense manifesteque atro maculatos, antheras 
sessiles, ad C. minutiftoro valde affinis sed ab ea petiolis canaliculatis (non planis) laminis 
abrupte largo- (non sub-) acuminatis, pedicellis 1 .5-2.5 (nee 0.5-1) mm longis, lobis calycinis 
secus margines hyalims (non opacis) serratis (nee crenulatis) punctato-lineatis (nee epunctatis) 
lobis corollinis secus marginis erosis (nee integerrimis) antheris acutis (nee ematginatis) 
facile cognoscitur. 

Treelet to 3 m tall. Branchlets angulate, 2-4 mm diam., densely rufous 
furfuraceous lepidote. Leaves alternate; blades chartaceous, elliptic to nar- 
rowly oblanceolate, (9-)l 1-15(-18) cm long, (3-)4-5(-6) cm wide, apically 
abruptly acuminate, basally cuneate, densely rufous lepidote (appearing granulose) 
above and below at first, glabrate above, somewhat persistent below, mid- 
rib impressed above, prominently raised below, the secondary veins 8-12 
pairs, impressed above, prominently raised below, inconspicuously pellu- 
cid punctate, the margin entire; petioles canaliculate, 1 .5-1 .8 cm long, glabrescent 
above, densely lepidote below. Staminate inflorescence a simple raceme, (4-) 
6_9(_10)cm long, the rachis and pedicels densely rufous furfuraceous lepi- 
dote; floral bracts carnose, linear lanceolate, 0.8-1.2 mm long, apically long- 
attenuate, the margin entire, densely lepidote; pedicels cylindrical, 1.9- 
2.2 mm long. Staminate flowers erect, 4-merous, carnose, subrotate, brownish-green; 

134 SiDA 18(1) 

calyx 0.8-1.1 mm long, the tube 0.1-0.2 mm long, the lobes ovate, 0.7- 
0.9 mm long, 0.8-0.8 mm wide, apically acute, abruptly constricted ba- 
sally, densely and prominently black punctate, moderately rufous lepidote, 
glabrescent, the margin hyaline, conspicuously black punctate-lineate, ir- 
regularly serrate, glabrous; corolla subrotate, 1.6-1.8 mm long, the tube 
0.3-0.4 mm long, square, glabrous, the lobes suborbicular, 1.3-1.4 mm 
long and wide, emargmate apically, abruptly constricted basally, densely 
and prominently black punctate, sparsely rufous lepidote without, glabrescent, 
densely glandular-granulose throughout within, the margin erose; anthers 
sessile at apex of corolla tube, thus appearing epipetalous, the tube 0.3-0.4 
mm long, glabrous, the anthers widely cuadrate, 0.3-0.4 mm long, 0.6- 
0.7 mm wide, apically acute, basally truncate, dehiscent by large subapi- 
cal, ovate pores, the pores not confluent, extending ca. 3/4 anther length, 
the connectives densely and prominently red or black punctate; pistillode 
conic, 0.2-0.3 mm long, sparsely glandular-lepidote. Pistillate inflomcence 
unknown. Fruit unknown. 

Distribution. — Endemic to the upper Rio Santiago Valley, in the Depart- 
ment of Amazonas, Peru, 180-200 m. 

Ecology and conservation status. — Cybianthus granulosus inhabits wet premontane 
forests above the Rfo Santiago valley, which together with the Rio Cenepa, 
comprise a region now known to be host to a number of endemic species. 
Given that it is most likely endemic, and not at all well-known, its conser- 
vation status cannot be determined at this time. 

Etymology. — The specific epithet refers to the densely rufous lepidote 
tomentum, which appears granulose when examined superficially. 

Par/viypes. PERU. Amazonas: Prov. Bagua, Dtto. Imaza, Comuniclad Aguaruna Putuim, 
Anexo Yamayakat, Zonas Alcas de Putuim, "Campou," 700 m, 18 Jan 1996 (Fr), C. Dun et 
al. 7649A (BRIT, HUT, MO, USM); Rio Santiago, 3 km from Caterpiza, 180 m, 1 2 Nov 
1 979 (scam, fl), V. H/iashikat 1221 (MO, USM); Valle del Rio Santiago, 65 km N of Pinglo, 
Quebrada Caterpiza, 2-3 km from Caterpiza, 200 m, 19 Sep 1977 (bud), V. H/mshikat677 
(MO, USM), 28 Nov 1979 (scam, fl), V. Huashikat 1422 (MO, USM). 

Cybianthus granulosus is most closely related to C. minutifiorus Mez by virtue 
of Its alternate, chartaceous leaf blades with acuminate apices and cuneate 
bases, densely rufous furfuraceotis-lepdidote branchles, flowers with carnose 
texture, densely and prominently black punctate, and sessile anthers. However, 
Cybianthus granulosus is easily separated from C. minuttjlorus by the abruptly 
long-acuminate leaf apices, canaliculate petioles, the hyaline, serrate, and 
punctate-lineate calyx margins, the erose corolla margins, acute anthers, and 
pedicels 1.5-2.5 mm long. 

35. Cybianthus flavovirens Pipoly, sp. nov. (Fig. 25). Typi;: PERU. San Martin: 
Prov. Mariscal Caceres, Dtto. Tocache Nuevo, Palo Blanco near Fundo de Manuel 

PiPOLY, Cybianthus in Ecuador and Peru 


Fic. 25. Cybianthus jlcivovirem Pipoly. A. Habit, sliowing anguiate branchlecs and gradually 
capering petioles. B. Abaxial leaf surface, showing sparse, rufous stellate tomentum. C. 
Section of inflorescence, showing nodding flowers, widely deltate calyx lobes and corolla 
with lobes constricted basally, prominent veins, and pusticulate surface. D. Branchlet, showing 
rufous stellate tomentum. A-D, drawn from holotype, by Linda Ellis. 

136 Sum 18(1) 

Aranjo, 700-800 m, 1 Mar 1979 (stam. fl)J, Sihimke 10883 (iiolotypi;: MO; isotypes; 

Propter ramulos angulatos, peciolos graciare angustatos, inflorcscentiam racemosam, flores 
nutanres, ancheras sessilia, C. imezi/elano valde arete affinis, sed ab ea lobis calycinis membranaceis 
(non carnosis) acucis (nee rotundatis) seciis margmis hyalinis (nee opaeis), antheris obtusis 
vel subacutis (non truncaris), porisque separatis (nee eonfluentibus), denique lobis eorollinis 
membranaceis (non carnosis) pustieulatis (nee planis) triineatis vel emarginatisque (nee aeutisque) 
praeclare distat. 

Tree to 4 m tall. Branchlets lightly angulate, 2.5-3.5 mm diam., densely 
rufous stellate-tomentose. Leaves alternate; blades chartaceous, elliptic, (12. 5-) 
19-.30(-32) cm long, (4-)6-10 cm wide, apically subacuminate to acumi- 
nate, the acumen 0.5-2 cm long, basally attenuate, dectirrent on the distal 
end of the petiole, midrib somewhat elevated but canaliculate above, not 
decurrent on petiole, prominently raised and densely rufous tomenose be- 
low, the secondary veins 10-16 pairs, dull green above, pallid green below, 
pellucid punctate above and below, sparsely rufous stellate-puberulent be- 
low, the margin entire, opaque, regular, flat; petioles semiterete (1.6-)2- 
2.5 cm, flat above, tapered, slightly thickend basally, not ptilvinate, gla- 
brous above, sparsely rufous pubescent below, glabrescent. Staminate infloresa')ice 
a lax raceme (7.5-)13-19(-40) cm long, the rachis and pedicels moderately 
rufous tomentellous, glabrescent; floral bracts linear-lanceolate, 1 . 1-1.5 mm 
long, 0.1-0.2 mm wide, apically attenuate, densely rufous tomentellous 
below, glabrescent, the margin entire; pedicels cylindrical, 2.5-3.5 mm long 
at anthesis, erect in bud, nodding in anthesis. Staminate flowers 4-merous, 
membranaceous, nodding, bright yellow-green; calyx cotyliform, 1.5-1.9 
mm long, the tube 0.6-0.8 mm long, the lobes widely deltate to triangu- 
lar, 1.1-1.3 mm long and wide, apically acute, epunctate, medially thick- 
ened, sparsely rufous puberulent, glabrescent, the margin hyaline, epunctate, 
entire, minutely glandular-ciliolate; corolla subrotate, 2.8-3 mm long, the 
tube 0.5-0.8 mm long, glabrous without, densely glandular-granulose within, 
the lobes suborbicular to oblate, 1.5-2.2 mm long, 2.2-2.6 mm wide, apically 
truncate to slightly emarginate, contracted basally, translucent, the three 
veins conspicuous, glabrous without, prominently pusticulate and sparsely 
glandular-granulose above but densely so toward base within, the margin 
opaque, sparsely glandular-granulose, entire; stamens apparently sessile at 
junction of corolla lobes and tube, the anthers sessile, very widely ovate- 
obcordate, 0.4-0.6 mm long, 0.5-0.8 mm wide, apically obtuse to sub- 
acute, basally cordate, the pores widely ovate, extending 1/2-3/4 anther 
length, separate (not confluent), the connective epunctate, minutely red glandular- 
papillate; pistiUode costate, subglobose, 0.8-1 mm long, 0.6-0.7 mm wide, 
hollow, the stigma capitate, densely yellow glandtilar-papilalte. Pistillate 
inflorescence: unknown. Fruit unknown. 

PiPOLY, Cybianthus in Ecuador and Peru 137 

Distribution. — Endemic to the junction of the westernmost Amazon Ba- 
sin (Hylaea) with the foothills of the Peruvian Andes, from 100—800 m elevation. 

Ecology and conservation status. — Cylnanthus flavovirens occurs in lowland 
moist forests on terra firme. These forests are drier than sites with the same 
general physiognomy farther to the north in Amazonas and Loreto. The few 
sporadic collections may be the result of underexploration rather than rar- 
ity, but owing to increasing pressure to cut forests for farming, this species 
should be considered threatened. 

Etymology. — The epithet refers to the bright yellow-green corolla, a unique 
feature in the subgenus. 

Paratypfs. PERU. Huanuco: Villa Isabel, Rfo Cuchara, 20 Sep 1961 (stam. fl),J. Schunke 
5667 (F, K, iVIO, US, USiVI). Loreto: Mishuyacu, near Iquitos, 100 m, Oct-Nov 1929 (scam, 
fl), G. Ktug 285 (F, US), (scam, fl), C. Kli/i^ 367 (F, US). Pasco: Prov. Oxapampa, Palcazu 
Valley, Cabeza de Mono, 5-6 km W oflscosaci'n, 10° 12' S, 75° I4' W, 14-15 Apr 1983 
(scam, fl), D. Smith 3 709 (MO, US, USM); Prov. Pasco, Palcazu Valley, Selva General, Cerro 
de Pasco, Proyccto Especial Pichis-Palcazu, IND, 09° 50' S, 68° OO'W, 300-600 m, 13 
Occ 1987 (stam. fl), G. Hartshorn i^t cil. 2996 (BRIT, MO, USM). 

Cybianthus flavovirens is a member of a complex of taxa related to C. venezuelanus 
Mez, as evidenced by the angulate branches, gradually tapered petioles, nodding 
flowers, and sessile anthers. However, C, flavovirens is easily recognized by 
its membranaceous perianth, acute calyx lobes with opaque margins, pusticulate, 
bright yellow-green corolla lobes with truncate or emarginate apices, and 
obtuse or subacute anthers with separate pores. This species has been con- 
fused with Cybianthus cyclopetalus. However, Cybianthus flavovirens may eas- 
ily be separated from that species by its angulate branchlets, racemose 
inflorescences, large, yellow flowers, and deltate to triangular calyx lobes. 

36. Cybianthus venezuelanus Mez in Engl., Pflanzenr. IV. 236(Heft 9):22T 
1902. Type: VENEZUELA. Lara: Barquisimeco, San Felipe, 600 m,Jun 1846 (scam, 
fl), N. Funck & L. SchliDi 678 (lhctotypf., here designaced: G-BOIS; isolectotypes: 
BM, BR,G,LD, P W). 

?eckia purpurea Rusby, Bull. New York Boc. Card. 4:405. 1907. Type: BOLIVIA. Wich- 
our localicy daca, (scam, fl), A.M. Bang 2048 (holotype: NY). 

Cybianthus egensis Mez in Engl., Pflanzenr. IV. 236(FIefc 9):222. 1902. syn. nov. Type: 
BRAZIL. Amazonas: Near Ega [Teffe], Sep 1831 (scam, fl), E. Poeppig 2567 (holo- 
type: W, F Neg. 31997; isotype: W) . 

Cybianthus hrownii Gleason, Bull. Torrey Boc. Club 53:293- 1926. syn nov. Type: GUYANA 
[BRITISH GUIANA]. Tumacumari, 18Jun-8 Jul 1921 (scam, fl), H. Gleason 159 
(holotype: NY; lsoi-ype,: K). 

Tree to 5 m tall. Branchlets angulate, (2.5— )3.5— 5 mm diam., densely ru- 
fous tomentose. Leaves pseudoverticillate; blades chartaceous, oblanceolate 
to elliptic, (l()-)17-27(-34)cm long, (3-)6-9(-ll) cm wide, apically acute 
to acuminate, basally acute, slightly decurrent on the petiole, midrib slightly 

138 SiDAlSd) 

raised above, prominently raised below, the secondary veins 10—13 pairs, 
rufous puberulent and smooth above, glabrescent, sparsely rufous puberu- 
lent below, concentrated along the midrib and secondary veins, inconspicuously 
pellucid punctate, the margin flat, entire; petioles slightly canaliculate, (15—) 
20— 25(— 30) mm long, tapered, densely and minutely stellate rufous stel- 
late puberulent. Staminate inflorescence: an erect, simple raceme, (7.5— )1 0.5— 
18(— 23) cm long, sparsely rufous stellate puberulent; peduncle 1—3 cm long; 
floral bracts coriaceous, lanceolate, 0.5-0.7 mm long, 0.2-0.3 mm wide, 
apically subulate, densely rufous stellate puberulent, the margin entire; pedicels 
cylindrical, 2.1—2.7 mm long, sparsely rufous stellate puberulent, glabres- 
cent. Staminate floicevs 4-merous, carnose, nodding, green; calyx cotyliform, 
0.9—1.1 mm long, the tube ca. 0. 1 mm long, the lobes widely ovate, O.cS— 
1 mm long, 0.9—1 mm wide, apically rounded, densely rufous stellate pu- 
berulent, sparsely and prominently black punctate, the margin opaque, coarsely 
serrulate, densely glandular-ciliolate; corolla carnose, subrotate, 1 .6—1.8 mm 
long, the tube quadrate, 0.5—0.6 mm long, glabrous, the lobes widely tri- 
angular, 1.1—1.3 mm long, 1.5—1.7 mm wide, reflexed in anthesis, apically 
actite, dorsally recurved, prominently and densely black punctate and gla- 
brous without, densely glandular granulose throughout within, the mar- 
gin slightly revolute, densely glandular-granulose; staminal tube inconspicuous, 
hyaline, membranous, 0.5—0.6 mm long, aclnate to corolla tube, anthers 
apparently sessile, cuadrate, 0.3-0.4 mm long, 0.5—0.6 mm wide, apically 
truncate, basally trtmcate, leaning proximally at anthesis, the connective 
prominently punctate dorsally, rufotis glandular-papillate apically; pistillode 
obsolete or conical, 0.4—0.5 mm long, 0. 1—0.2 mm wide, hollow, glabrous. 
Pistillate inflorescence as in staminate but (4.5— )8.5-l ()(-l 3) cm long; peduncle 
1—2.5 cm long; floral bracts 0.5—0.7 mm long, 0.2—0.3 mm wide; pedicels 
0.9—1.2 mm long. Pistillate flowers as in staminate but calyx 0.8—1.2 mm 
long, the ttibe ca. 0.2 mm long, the lobes 0.6—0.8 mm long, 0.9— 1-1 mm 
wide; corolla 1.2—1 .4 mm long, the ttibe 0.4-0.5 mm long, the lobes 0.7— 
0.9 mm long, 1.1 — 1.2 mm wide; staminodial as in staminal tube, 0.4—0.5 
mm long, adnate to corolla tube, the antherodes 0.2—0.3 mm long, 0.4— 
0.5 mm wide; pistil obturbinate, 0.5—0.6 mm long, 0.3—0.4 mm wide, glabrotis, 
the ovtiles 2—3, partially imbedded in the placenta. Fruit globose, 5— 7(— 9) 
mm diam., black at maturity, the exocarp thick, juicy. 

Distribution. — In moist forests rimming the Amazon Basin, from Guyana 
through Veneztiela to the Andes of Colombia sotithward to Bolivia, 100— 
1 ,00{)(-2,000) m. It is also known from the Choco Floristic Province of Colombia, 
and may be expected in that corresponding region of Ecuador. The Boliv- 
ian specimens cited below represent new distribution records. 

Ecology and conservation status. — Cyhianthus venezuelanus is common in lowland 
and premontane moist forests, and occasionally in premontane pluvial for- 

PiPOLY, Cybianthus in Ecuador and Peru 139 

ests. Ir is locally common and appears to do well in light gaps and forest 
margins, thus, it should not be considered threatened. 

Etymology. — The epithet refers to the the country from which the type 
was collected, Venezuela. 

Representative specimens examined. ECUADOR. Zamora-Chinchipe: Nangaritza Canton, 
Pachicutza, Camino al Hito, Cordillera del Condor, 04" 07' S, 78° 37' W, 1,000-1,100 m, 
19 Oct 1991 (fr), W. Palachs et al. 8313 (BRIT, MO, QCNE). PERU. Huanuco: Prov. 
Leoncio Prado, Dtto. Rupa Rupa, Ynti, Rd. to Ri'o Rondos, 750 m, 24 Mar 1972 (stam. 
fl),./. Sc/y/inke 3308 (AMAZ, F, G, MO, NY, US, USM). Loreto: Prov Maynas, Mishuyacu, 
near Iquitos, 100 m, Oct-Nov 1929 (stam. fl), G. Ki//g 94 (F, NY, US); Peria Negra, near 
Iquitos, 100 m, 20 Oct 1979 (bud), R Ayaia2I02 (AMAZ, MO, USM); QuebradaOrejon, 
Purma, 29 Oct 1980 (fr), F. Aya/a et al. 2814 (AMAZ, MO, US, USM); Pumayacu, be- 
tween Balsapuerto and Moyobamba, 600-1 ,200 m, Aug-Sep 1 933 (stam. fl), G. Kh/s^ 3188 
(F, G, MO, NY, S, US). Madre de Dios: Prov. Tambopata, Tambopata Wildlife Reserve, 
30 km S of Puerto Maldonado, 12° 15' S, 69° 17' W, 260 m, 14 Nov 1984 (stam. fl), H. 
Young et al. 1 79 (MO, US, USM); Tambopata Tourist Camp at junction of Rfos Tambopata 
and La Torre, 12° 49' S, 69° 43' W, 280 m, 22 Jul 1985 (ster.), A. Gentry et al. 51083 
(CUZ, MO, USM), 27 Jul 1985 (ster.), A. Genttyetal 5 1400 (CUZ, MO, US, USM). BOLIVIA. 
Santa Cruz: Prov. Florida, Parque Nacional Amboro, Sta. Rosa de Lima, 5-8 km N of 
Cerca a La Cumbre, on path to La Playa, 1 7° 49.5' S, 64° 16' W, 2,000-2,100 m, 3-5 May 
1993 (fr), /. Vargas et al. 2248 (BRIT, MO, USZ) 

The perianth of the lectotype o( Cybianthus venezuelanus , with dense and 
prominent black punctations, the truncate sessile anthers, large oblanceolate 
chartaceoLis leaves, and short pedicels match the collections cited here. Cybianthus 
venezuelanus , as treated here, is a polymorphic ochlospecies with many re- 
gional variants. The type q{ Cybianthus brownii Gleason varies only in its 
shorter pedicels and longer corolla lobes. Likewise, the type o{ Cybianthus 
egensis Mez, in bud, has correspondingly shorter pedicels and corollas. Cybianthus 
venezuelanus is easily recognized by the angulate branchlets, pseudoverticillate 
leaves, rounded calyx lobes with opaque margins, and truncate anthers with 
confluent pores. Its distribution, which rims the Amazon Basin, is unusual 
in the Myrsinaceae but is common in other families, such as the Piperaceae 
(R. Callejas, pers. comm.). 

37. Cybianthus grandezii Pipoly, sp. nov. (Fig. 26). Typh: PERU. 
Prov. Maynas, Quebrada Yanayacu, entering from Aucayo, 25 Aug 1990 (stam. fl), 
C. Grdndez. S. Vdsquez & M. F lores 1824 (holotype: MO; isotypics: AMAZ, US). 

Quoad folia magna chartacea nervos secundarios tertiarioscjue laminares praeclare utrinque 
conspicLia ad bases gradatim descrescentiaque necnon petiolos pulvinatos C. jensoni valde 
affinis sed ab ea laminis ad bases obtusis auriculatisque (non cuneatis) manifeste desu|"ier 
perpuncticulosis (nee epunctatis) desuper nitidus (nee sordidis) nerviis secundariis 18-24 
(non 24-30) -jugis, 12-13.5 (non 6.5-9) cm latis, petiolis 0.5-1.4 (non 2. 1-5) cm longis, 
pedicellis 0.3-0.5 (non 0.8-1.4) mm longis, bracteis florinis pedicellis 3 (non 6-7)-plo 
longiores statim cognoscitur. 

Shrub to 2 m tall. Branchlets terete, ca. 10 mm diam., rufous-lepidote. 


SiDA l.Sd) 

Fk ;. 2fi. Cyhianthi/s graiulezn Pjpoly. A. Habit, showing basally cruncace and somewhat auricukite 
leaf bases. B. Adaxial, prominently perpuncticuiose surface. C. Abaxial, minutely rutous 
lepidote surface. D. Portion ot inflorescence. E. Pistillate calyx, showing pustulate, black 
punctate lobes with hyaline, erose-serrulate margins. F. Pistillate cotolla, showing pustu- 
late abaxial surface. G. Branchlet apex. A— Ci, drawn from holotype by Linda Ellis. 

Pipoi Y, Cybianthiis in Ecuador and Peru l4l 

Leaves ahernate; blades chartaceous, oblanceolate, 34—36 cm long, 12—13-2 
cm wide, apically subacuminate, basally truncate and appearing auriculate, 
nitid above, pallid below, glabrous, and prominently black perpuncticulose 
above, sparsely and minutely rufous-lepidote below, the midrib raised and 
canaliculate above, prominently raised, black punctate-lineate and densely 
rufous-lepidote below, the secondary vems 18-24 pairs, loop-connected 
submarginally, the margin entire; petioles canaliculate, (0.5-)l-1.4 mm 
long, 0.4—0.6 mm diam., pulvinate, densely lepidote. Staminate inflorescence 
unknown. Pistillate inflorescence an erect, straight simple raceme, (8.5— )1 1— 
14 cm long; peduncle 1-3 cm long; the rachis green, densely red glandu- 
lar-papillate, black punctate-lineate; floral bracts membranaceous, linear- 
lanceolate, 1.8-2 mm long, densely rufous lepidote; pedicels obconic, 0.5-1 
mm long, densely glandular-papillate. Pistillate flowers chartaceous, creamish- 
white; calyx cotyliform, 1.8-2 mm long, the tube ca. 0.5 mm long, the 
lobes widely ovate, 1.3-1.5 mm long, 1.1-1.2 mm wide, apically acute, 
densely and prominently black punctate, prominently translucent pustu- 
late, the margin hyaline, irregular, erose-serrulate, epunctate, glabrous; corolla 
subrotate, 2.2-2.4 mm long, the tube ca. 0.5 mm long, the lobes very widely 
ovate 1.7-1.9 mm long and wide, apically rounded, densely and promi- 
nently black punctate and translucent pustulate without, glandular-granulose 
throughout and prominently black punctate within, the margin hyaline, 
epunctate, minutely erose- crenulate, glabrous; staminodes 1.1-1.2 mm long, 
staminal tube conspicuous, carnose, 0.4-0.5 mm long, elobate, densely glandular- 
papillate, the filaments terete, proximally curved, 0.2-0.3 mm long, the 
sterile anthers ovate, 0.5-0.6 mm long, and wide, the apically apiculate, 
dehiscent by terminal confluent pores ca. 3/4 length, the connective prominently 
black punctate; pistil obturbinate, 1.5-1.6 mm long, 0.5-0.6 mm diam., 
densely and prominently black punctate, translucent glandular-lepidote, 
the ovary 0.5-0.6 mm long, the style 0.8-0.9 mm long, the stigma bi- 
lobed, the lobes to 0.1 mm long, distally curved, the placenta subglobose, 
with 2-4 ovules partially embedded. Fruit globose, 8-10 mm long and in 
diam., the exocarp thick, juicy, purple-black at maturity. 

Distribution. — Known only from Maynas Province, Dept. of Loreto, Peru, 
at up to 125 m elevation; presumably endemic. 

Ecology and conservation status. — Cybianthiis grandezii is restricted to pri- 
mary lowland most upland terra firme forest over red lateritic clays. Label 
data indicate it is rare, and given the valuable timber present in that forest 
type, this species should be considered threatened. 

Etymology. — It is indeed a pleasure to dedicate this species to Biol. Cesar Grandez, 
professor of biology at the Universidad Nacional de la Amazonia Peruana 
(UNAP), Iquitos, and authority on the systematics of Peruvian Flacourtiaceae. 
Cesar is an ardent field worker, an excellent teacher and herbarium curator. 

142 SiDA 18(1) 

Para'iypf,: PERU. Loreto: Prov. Maynas, Drto. Fernando Lores, Panguana, 125 m, 6 
Aug 1 991 (fr), S. AlcDc/i/n-I & M. R/madn 31219 (MO, IBE). 

Cyhianthus grandezii is closely related to C. jensonii Pipoly, but is easily 
recognized by its subsessile leaves with truncate bases. The perpuncticulose 
and nitid adaxial leaf surface, and secondary veins raised prominently above 
and below are also distinctive. 

38. Cybianthus jensonii Pipoly, sp. nov. (Fig. 27). Typi;: PERU. Loreto: Prov. 
Alto Amazonas: Andoa.s, Rio Pastaza near Ecuadorean border, 02° 48' S, 76° 28' W, 
210 m, 14 Aug 1980 (fr), A. Gentry. R. Vasqi/ez & N. Jaramillo 29700 (holotyph: 
MO; isoTYPns: AMAZ, NY, USM). 

Quoad lolia magna charracea nervos laminare.s secundarios tertianosque praeclare utrinque 
conspicLia ad bases gradacim aqdescrescentia i^eciolosque pulvinatos C. grandezi valde affinis sed 
ab ea laminis ad bases cuneatis (nee obtusis auricularisque) epunccatis (non manifeste desuper 
perpuncticulosis) desuper sordidis (non nitidis) nerviis secundariis 24-.^0 (non 18-24)- 
JLigis, 6.5-9 (nee 12-1 3.5) cm latis, pctiolis 2.1-5 (non 0.5-1 .4) cm longis, pedicellis 0.8-1.4 
(non 0..^-().5) mm longis bracteis florinis pedicellis 6-7 (non 3)-lilo longiores statim cognoscitur. 

Treelet to 4 m tall. Branchlets terete, 7-9 mm diam., lenticellate, the bark 
vertically ridged, densely rufous tomentose. Leai'es alternate; blades mem- 
branaceous, oblanceolate, .33^9 cm long, 6.5—9 cm wide, apically subacuminate, 
basally long-attenuate, dull green above and below, the veins 24-30 pairs, 
buUate above, prominently raised below, the tertiary areoles prominently 
raised below; glabrate above, moderately rubiginous furfuraceous lepidote 
and sparsely pellucid punctate below, the margin entire, decurrent, gradu- 
ally tapering to the petiole; petioles marginate, 2.1-5 cm long, to 0.5 cm 
thick, glabrous, abruptly pulvinate, the pulvinus 1.3-2 cm above petiole 
base. Staminate inflorescence: a simple, lax, axillary raceme 6.5—8 cm in bud; 
the rachis, bracts and pedicels densely rufous tomentose; floral bracts mem- 
branaceous, linear-lanceolate, 3^ mm long, 0.3-0.4 mm wide, apically subulate, 
the margin entire; pedicels cylindrical, 0.8-1 .4 mm long, glabrescent. Staminate 
flowers 4-merous; calyx membranaceous, cotyliform, 1.4-1.6 mm long, the 
tube 0.2-0.4 mm long, the lobes widely ovate, 1-1.3 mm long, 1.2-1.5 
mm wide, apically acuminate, densely and prominently black punctate, sparsely 
rufous pubescent, the margin hyaline, flat, erose, epunctate; corolla (in bud) 
subrotate, membranaceous, to 1.2-1.5 mm long, the lobes ovate, apically 
obtuse, densely and prominently black punctate, glabrous without, sparsely 
glandular-granulose within, the margin hyaline, flat, erose, epunctate, gla- 
brous; stamens appearing epipetalous, the anthers sessile at the junction of 
corolla tube and lobe, deltoid, ca. 0.7 mm long and wide, dehiscent by subapical 
pores, the connectives red punctate medially; pistiUode, conic, hollow, ca. 
0.5 mm long and 0.2 mm diam. Pistillate inflorescence as in staminate but 3- 
6(-7.2) cm long; floral bracts 1-1.6 mm long, 0.2-0.4 mm wide, apically 
long-attenuate; fruiting pedicels incrassate, 0.7-0.9 mm long, to 1.5 mm 

PiPOi.Y, Cybianthus in Ecuador and Peru 


Fic. 27. Cybtanthiis jemunit Pipoly. A. Habit, showing long-attenuate leat bases. B. Abaxial 
leaf surface with prominently defined areoles. C. Portion of staminate inflorescence in bud. 

D. Open staminate bud. E. Pedicel, calyx and fruit. A-D, drawn from Gentry et al. 33708. 

E, drawn from Croat 19485. Figure drawn by Linda Ellis. 

144 SiDA 18(1) 

diam. Pistillate flowers as in staminate, calyx 1.4-1.6 mm long, the tube 
0.2-0.4 mm long, the lobes 1-1.3 mm long, 1.2-1.5 mm wide; corolla, 
staminodes and pistil unknown. Fr//// depressed-globose, orange, 5—6 mm 
long, 6-7 mm diam., smooth, prommently pellucid punctate. 

Distribution. — Endemic to lateritic slopes above riparian areas in the 
Department of Loreto, Provinces of Alto Amazonas and Maynas, in the northern 
Amazon Basin of Peru, 130-210 m. 

Ecology and conservation status. — This species occurs in primary lowland 
tropical varzea forest margin. It is surely a rare species, given the recent 
massive collection effort at the sites during the Florula of the Biological Re- 
serves of Iquitos project (Vasquez 1997), during which the species was not 
relocated at the Explorama Inn (Indiana) site. Given its rarity, this species 
should be considered threatened. 

Etymology. — I dedicate this species to Peter Jenson, President of Explorama 
Tours, conservationist and principal promoter of ecotourism in the Peru- 
vian Amazon. Much of our knowledge regarding the biology of tropical 
ecosystems in the Peruvian Amazon would not have been discovered were 
it not for Peter and his associates' enthusiastic support, generosity and hos- 
pitality at the company's biological reserves (Explorama Inn, Explorama Lodge, 
and Explornapo Camp) which now serve as long-term ecological study sites. 
Explorama Tours' properties, with their combination of research, public education 
and collaboration with local communities and tourism, serves as the most 
successful model tor tropical ecotourism known. 

Parat^'pes. PERU. Loreto: Prov. Maynas, Explorama Inn, 2 km W of Indiana on Rio 
Amazonas, 03° 30' S, 73° 02' W, 130 m, 12 Feb 1987 (scam, bud), A. Gentry et ^i I. 55708 
(AMAZ, MO); Varadero de Mazan from Rio Amazonas to Rfo Napo, 22 Aug 1972 (fr), T. 
Cnuit 19485 (AMAZ, MO, NY, USM). 

Cybianthus jensonii is most closely related to C. grandezii Pipoly, but eas- 
ily recognized by its long-attenuate leaf bases, epunctate, pallid abaxial leaf 
surfaces, the secondary veins 24—30 pairs, narrower leaves with much longer 
petioles, subobsolete pedicels 0.8—1.4 mm long, and longer floral bracts. 

39. Cybianthus fosteri Pipoly, sp. nov. (Fig. 28). Type: PERU. Madre de Digs: 
Prov. Manu, Atalaya, vicinity Hacienda Amazonia, 2-3 km W of village, across Rio 
Alto Madre, 12'^^ 55' S, 71° 12' W, forested ridge, 600-900 m, 7 Dec 1983 (stam. 
fl), R. Foster & T. Wachter 7254 (iiolotype: MO; isotypes: F, NY, USM). 

Ob folia pseudoverticillata magna charcacea longipetiolata abrupte acuminataque, inflorescentia 
longiracemosa, flores nutantes, coriaceosc]ue, antheras sessiles manifesce necnon dorso punctatas, 
C. venez/zelano valde affinis sed ab ea ramulis teretes (non angulacis), foliis pscudoverticillatis 
(non alternis), laminis denseque manifeste atro-punctatis et omnino prominens (non parceqe 
plane subter atro lineato-punctatis) petiolatis canaliculatis (non marginatis ad bases abrupte 
crassis (nee gracilis) lobis corollinis interius pustulatis (non planis), antheris ad apices rocundatis 
(non truncatis) poris separatis (non conHuentibus) praeclare distat. 

PiPOLY, Cybiantlius in Ecuador and Peru 


Fig. 28. Cyhianthus fosten Pipoly. A. Habit, showing terete branchlets and pseudoverticillate 
phyllotaxy. B. Portion of staminate inflorescence, showing pustulate corolla and apicalUy 
rounded anthers with separate (not birimose) pores. C. Abaxial leaf surface detail, showing 
punctations and sparse puberulence. D. Branchlet apex, showing tomentum. A-D, drawn 
from holotype, by Linda Ellis. 

146 SiDA 18(1) 

Tree to 5 m tall. Branchlets terete, (6-)7-9 mm diam., densely terrugineotis 
tomentose. Leaves pseudoverticiUate; blades chartaceous, oblanceolate, (26-) 
28—34 cm long, 5.5-9(-H)) cm wide, apically abruptly acuminate, basally 
long attenuate, midrib somewliat elevated above, prominently raised be- 
low, the secondary veins 11 — 18 pairs, dull green above, pallid below, prominently 
black punctate above and below, sparsely rufous puberulent below, the margin 
entire, regular, flat; petioles canaliculate, 1 .6—2 cm, pulvinate, ferrugineous 
tomentose, glabrescent. Stanihiate ijiftorescence a lax raceme 8—12 cm long, 
moderately rufous lepidote, glabrescent; floral bracts lanceolate, 2—3 mm 
long, 0.8—1 . 1 mm wide, apically acute, densely rufous lepidote; pedicels cylin- 
drical, 3.5—5 mm long at anthesis erect in bud, nodding in anthesis, densely 
rufous lepidote. Stcoiiinate flowers 4-merous, coriaceous, nodding, pale green; 
calyx cotyliform, 1.2—1.3 mm long, the tube 0.5—0.6 mm long, the lobes 
widely ovate, 0.7— 0.8 mm long, 1.1 — 1 .2 mm wide, apically rounded, densely 
and prominently black punctate, sparsely rufous lepidote, glabrescent, the 
margin stramineous, opaque, epunctate, erose-fimbriate, glabrotis; corolla 
subrotate, 2-2.5 mm long, the tube 0.5—0.6 mm long, glabrous, the lobes 
widely ovate, 2—2.1 mm long, 1.4—1.6 mm wide, apically acute, densely 
and proniinently black punctate, sparsely rtilous lepidote without, glabrescent, 
prominently pustulate and densely glandular-granulose throughout within, 
the margin stramineous, erose, glandular-granulose; stamens apparently sessile 
at junction of corolla lobes and tube, the anthers sessile, very widely ovate, 
0.5—0.6 mm long, 0.8—0.9 mm wide, apically rounded, basally truncate, 
the pores widely ovate, extending 1/2—3/4 anther length, separate (not conl^tient), 
the connective prominently red ptmctate; pistiUode subglobose, 0.7-0.8 
mm long, 0.3—0.4 mm wide, hollow, densely glandular-lepidote. Pistillate 
inflorescence as in staminate but erect, 3—5 cm long; floral bracts 2—3 mm 
long, 0.8—1.1 mm wide; pedicels 2.5—4 mm long in fruit. Pistillate flowers 
as in staminate but calyx 0.8—1 mm long, the tube 0.2—0.3 mm long, the 
lobes 0.5—0.7 mm long, 0.9—1.1 mm wide; corolla, staminodes and pistil 
unknown. Fr/zit globose, 6—8 mm long and diam., red at maturity. 

Distribution. — Known only from the type locality, presimiably endemic. 

Ecology and conservation status. — Cyhianthi/s fosteri is endemic to one area 
of the Manti Biosphere Reserve and National Park, one of the largest in 
South America. It is a ridgetop species in lowland topical most forest. Given 
the extension of the Reserve and the species narrow range, it should not be 
considered threatened at this time. 

Etymology. — This species is dedicated to Robin Foster, of the Smithsonian 
Tropical Research Institute and a research associate of the Field iVIuseum of 
Natural Flistory. Robin has served as one of the co-founders of the Rapid 
Assessment Protocol, and is an expert on the reproductive biology oiTacbigali 
(including Sclerolohiitin) of the Fabaceae. 

PiPOi,Y, Cybianchus in Ecuador and Peru 147 

Paratyfe: PERU. Madre de Dios: Prov. Manii, Aralaya, vicinity Hacienda Amazonia, 
2-3 km W of village, across Rfo Alto Madre, 12° 55' S, 71° 12' W, 6()()-90() m, 7 Dec 
1 983 (fr), R. Foster^G T. Wachter 7242 (BRIT, F, MO, NY, USM). 

Cybianth//s fosteri is closely related to Cybianthus venez/ielanus , but is eas- 
ily recognized by its terete branchlets, pseudoverticillate leaves prominently 
and densely black punctate above and below, canaliculate petioles, pustu- 
late corolla lobes, and rounded anthers with separate (not birimose) pores. 
The pustulate corolla lobes are unique within the genus. 

40. Cybianthus resinosus Mez in Engl., Pflanzenr. IV. 236(Heft 9):219. 
1902. Typh: PERU. Loreto: Prov. Maynas, near Yurimaguas, without date (fr), E. 
Poejipig 2428 (holotype: W; isotypc: P). 

Tree to 1 5 m tall. Branchlets 2.5—3.5 mm diam., densely ferrugineous dendroid- 
tomentose at first, glabrescent. Leaves pseudoverticillate; blades thinly co- 
riaceous, elliptic, (ll-)15-21(-26) cm long, (4-)5.5~7(-l().8) cm wide, 
apicaliy long-acuminate, the acumen 0.8— 1.5(— 2) cm long, basally cuneate, 
decurrent on the petiole, midrib prominently elevated above and below, 
decurrent to petiole base, the secondary veins (9-)l 1-16(-18) pairs, nitid 
and glabrous above and below, inconspicuously pellucid-punctate (not vis- 
ible when dried), the margin entire, irregular, flat, entire; petioles canaliculate, 
l-2(-3) cm long, tapered, densely ferrugineous dendroid-tomentose, gla- 
brescent. Staminate inflorescence a lax, simple raceme, 1 1-1 5 cm long, sparsely 
rufous stellate puberulent; peduncle 0.9-l(-l-8) cm long; floral bracts 
chartaceous, linear, 1—1.2 mm long, 0.2—0.3 mm wide, apicaliy subulate, 
densely rufous stellate puberulent above and below, the margin entire, early 
caducous; pedicels cylindrical, 2.1-1.7 mm long, sparsely puberulent, gla- 
brescent. Staminate flowers 4-merous, membranaceous, nodding, greyish-brown; 
calyx cotyliform, 0.9-1 mm long, the tube 0.1-0.2 mm long, the lobes 
widely ovate, 0.6-0.8 mm long, 0.9—1 -2 mm wide, apicaliy acuminate, sparsely 
rufous stellate puberulent, densely and prominently black punctate, the margin 
hyaline, erose, short glandular-ciliate; corolla subrotate, translucent, 1.6- 
1 .8 mm long, the tube cylindrical, 0.3—0.5 mm long, the lobes very widely 
ovate, 1.2-1.5 mm long, 1.5-1.8 mm wide, flat, apicaliy obtuse, densely 
and prominently black punctate and glabrous without, densely glandular- 
granulose and pusticulate within, the margin irregular, glandular-granulose, 
entire; stamens 0.7—0.9 mm long, the tube inconspicuous, membranaceous, 
0.3-0.5 mm long, sessile, the anthers cuadrate, 0.2-0.3 mm long, 0.5-0.6 
mm wide, apicaliy truncate, translucent, glabrous, the connective promi- 
nently red punctate dorsally; pistillode cylindrical, 0.3-0.5 mm long, 0.2- 
0.3 mm diam., hollow, densely punctate, glabrous, the stigma 3-lobed. Pistillate 
inflorescence -AS in staminate but (l-)1.5-5 cm long; peduncle (0.3-)O.5-l(- 
1.5) cm long; floral bracts 0.6-1 mm long, 0. 1-0.2 mm wide; pedicels slightly 

148 SiDAl8(l) 

obconical, 0.7—1. 1 mm long, erect in fruit. Pistillate flowers as in staminate 
but 1-1.2 mm long, the tube 0.2— 0.3 mm long, the lobes 0.8-0.9 mm 
long, 1-1.2 mm wide; corolla 1.3-1.5 mm long, the tube 0.3—0.5 mm long, 
the lobes 0.8—1 mm long, 1.1—1.5 mm wide; stammodes as in stamens but 
0.4-0.6 mm long, the tube 0.3-0.5 mm long, the antherodes ca. 0.1 mm 
long, 0.2—0.3 mm wide; pistil obturbinate, 0.4—0.6 mm long, and in di- 
ameter, the style very short, the stigma 3-lobed, the placenta cotyliform, 
bearing 2 apically exposed ovules. Fr/iit globose, 5—7 mm diam. at matu- 
rity, exocarp black, juicy, edible at maturity. Bisexual inflorescence ^s in pis- 
tillate but a lax, simple raceme, or rarely a poorly formed panicle, 5-8(— 1 0) 
cm long; pedtmcle 0.5-1 cm long; floral bracts 1-1.2 mm long, 0.2-0.3 
mm wide; pedicels 1.2—1.5 mm long. Bisexual flowers as in pistillate, but 
stamens as in staminate, 0.5-0.8 mm long, the tube 0.2-0.3 mm long, the 
anthers ca. 0.3—0.5 mm long, 0.4—0.5 mm wide; pistil as in pistillate, conical, 
0.4-0.6 mm long, and in diam. Bisexual jriiit globose, A~G mm diam. at 
maturity, exocarp reddish-black, thin. 

Distribution. — Once thought to be endemic to the Iquitos area of Loreto, 
Peru, Cybianthus resinosus is now known (reported for the first time here), 
from the Choco of Colombia, Amazonian Ecuador, Venezuela, with one disjunct 
population in French Guiana {Oldeman 5272) growing at 1 00-200(-l ,300) 
m elevation. 

Ecology and conservation status. — Cybianthus resinosus is restricted to pri- 
mary non-inundated forests on white sand (varillal). While it is locally abundant, 
it should be considered threatened due to incre-asing habitat loss. The Ecuadorean 
and Venezuelan populations are unusual because they occur in premontane 
pluvial forest and wet forest on lower tepui talus slopes (on sandstone) re- 
spectively, each containing numerous lowland elements. It may be expected 
in the Rio Cenepa-Rio Santiago Drainage Basins, of Amazonas, Peru, an 
area known to show the same environments with numerous pockets of sandstone. 

Etymology. — The specific epithet refers to the highly nitid adaxial leaf surface, 
giving it a laccjuercd, resinous appearance. 

Specimens examined. COLOMBIA. Valle del Cauca: Bajo C;alima Concession, ca. 2") 
km NW of Buenaventura, 9 km NW of San Isidro intersection on "Canalete," near gate, 
5-45° slopes, 03° 59' N, 77" 08' W, 50 m, 13 Jul 1988 (ster.), D. Fciher-Langendoen &J. 
Hurtado 1 757 (CUVC, MO). VENEZUELA. Territorio Federal Amazonas: Dept. Atabapo, 
base of cliff and forest below slope of Cerro Huachamacari, 03" 39' N, 65° 43' W, 1 ,000- 
1,300 m, 5 Mar 1985 (stam. 11), R. Liesiier 18302 (BRIT, MO, VEN). ECUADOR. Napo: 
Canton El Chaco, Rfo CiranadiUo, Campamento de INECEL, "Codo Alto," 00" 08' S, 77" 
28' W, 1,300 m, 13-15 Sep 1990 (fr), W. Pa/acm 5389 (MO, QCNE). PERU. Huanuco: 
Prov. Pachitea, region of Pucailpa, W part of Sira Motintains and adjacent lowland, ca. 24 
km SE to 26 km ESE of Puerro Inca, from Campamento Sira, 09" 28' S, 74° 17' W, SE to 
valley of Rfo Negro, 750 m, 29 May 1988 (fr), B. Wal/mfcr 14-29388 (BRIT, MO, W, 
WU). Loreto: Prov. Maynas, Iquitos, May 1925 (stam. fi), G. Tessiiu/nii 3143 (NY); Casen'a 

PiPOLY, Cybianthus in Ecuador and Peru 149 

Mishana, 30 km SW of Iquitos, Callicebus Biological Reserve, 4 km S of Mishana, 19 Aug 
1980 (stam. fl), R. Foster 4404 (F-2 sheets, NY); Mishana, l6 Aug. 1978 (pist. fl, h),J. 
Ramtrez 17 (AMAZ, MO); Mishana, Rfo Nanay, 03° 50' S, 73° 30' W, 140 m, 16 Aug 
1978 (fr),./. Ramiyez 132 (AMAZ, MO); Mishana, along Rio Nanay, 03° 55' S, 73° 35' W, 
150 m, 20 Jan 1985 (fr), R. Va'squez & N. Jaramillo 6126 (AMAZ, MO, NY); AUpahuayo, 
Estacion IIAP, 04° 10' S, 73° 30' W, 150 m, 5 Jun 1985 (bud), R. Va'squez et al. 6551 
(AMAZ, BRIT, MO, NY), 29 May 1990 (fr) R. Vdscjuez et al. 13764 (AMAZ, MO, USM), 
16 Aug 1990 (pist. fl, fr), R. VascjM'z & H. Jaramillo 14224 (AMAZ, BRIT, F, MO, NY, 
TEX, US, USM), 4 Dec 1990 (fr), R. Va'squez & N. Jaramillo 15237 (AMAZ, BRIT, E, MO, 
US, USM), 150-180 m, 29 May 1991 (ster.), R. Va'squez & N. Jaramillo 166H1 (AMAZ, 
BRIT, MO, USM), 150 m, 23 Mar 1992 (ster.), R. Vasquez et al. 17996 (AMAZ, BRIT, 
MO, USM); AUpahuayo, ca. 26 km along Iquitos-Nauta Rd., 130 m, 25 Aug 1988 (fr), H. 
van cler Werff 10273 (AMAZ, MO); Mishuyacu, near Iquitos, 100 m, Sep 24-28 1929 (fr), 
E. Ktllip & A. Smith 29873 (E, US), May-Jun 1930 (bisex. fl), G. Klug 1384 (F, NY, US); 
Quistococha, 200 m, 27 May 1978 (pist. fl, fr), A. Gentry & N. Jaramillo 22314 (AMAZ, 
MO); Altura de Pina Negra, SW of Iquitos, ca. 3-4 km past Quistococha, 200 m, 19 Nov 
1975 (fr); Caseri'o de Urcumirafio, Rfo Napo, path from settlement to tall foresr, 120 m, 8 
Oct 1979 (fr), C. Diaz ON. Jaramillo i486 (MO, NY); Pena Blanca, on Rfo Itaya, 110 m, 
19 Sep 1929 (fr), E. Ktllip & A. Smith 29(>72 (E, US); Between Yurimaguas and Balsapuerto 
(lower Rfo Huallaga basin), 1 35-1 50 m, 26-31 Aug 1929(fr)£. Ktllip & A. Smith 28110 
(F, NY, US); Prov. Requena, Dtto. Sapuena, Jenaro Herrera, Rfo Ucayali, 04° 55' S, 73° 
40' W, 160 m, 16 Aug 1994 (stam. fl), R. Ortiz et al. 101 (AMAZ, BRIT, MO); without 
locality, except "in Peruvia subandina, without date (fr), E. Poeppig s.n. (L). 

Cybianthus resinosus is most closely related to C. penduliflorns Mart., but is 
easily separated from it by the inconspicuously punctate leaves, longer pedicels, 
flat corolla lobes and calyx lobes with acuminate apices and erose, short- 
ciliate margins. The adaxial prominently raised midrib decurrent to the petiole 
base is unique within the subgenus. The fruit oi Cybianthus resinosus is also 
smaller, black and has a thick exocarp, and it inhabits terra firme forests on 
white sand whereas C. penduliflorus is an igapo species. 

41. Cybianthus fuscus Mart., Flora 259- 1841. Typr: BRAZIL. MatoGrosso: 
"Prope rivum Cochim in Cujaba," May (pist. fl), P. da Silva Manso s. n. (i iolotyfi;: M) 

Shrub or small tree to .3 m tall. Branchlets terete, 2-.3 mm diam., densely 
dendroid and stellate rufous glandular-tomentose, tardily glabrescent. Leaves 
pseudoverticiUate; blades chartaceous to coriaceous, very narrowly oblan- 
ceolate or very narrowly oblong, (13-)l6-25(-30) cm long, 2-4(-6) cm 
wide, apically long acuminate-attenuate, the attenuated portion 1-2 cm 
long, terminating in a minute rounded tip, basally long acuminate-attenu- 
ate, the attenuated portion 1.5-2 cm long, giving the petioles appearance 
of being longer, fully decurrent on petiole to pulvinus; midrib prominently 
elevated above, decurrent to petiole base, the secondary veins 12-25 pairs, 
somewhat to deeply impressed, the leaf appearing subbullate to buUate above, 
prominently raised and loop-connected below, somewhat nitid and glabrous 
above, pallid, rufous papillate and conspicuously black punctate and punc- 
tate-lineate below, the margin entire, flat, glabrous; petioles somewhat marginate, 

1^'* SiDA 18(1) 

5-l()(-12) mm lon^ij;, with a basal pulvinus. Staminate inflorescence: a lax raceme 
(2.5-)5-8(-19) cm long; peduncle 0.8-1 .5 cm long; floral bracts chartaceotis, 
linear, 1 .2-1.5 mm long, 0.3 mm wide, apically subidare, densely and promi- 
nently rufous papillate; pedicels 3.5-5 mm long, densely papillate . Stami- 
nate flowers chartaceous, 4-merous; calyx cotyliform, 0.8-l(-l .8) mm long, 
the tube ca. 0.2 mm long, the lobes very broadly ovate or linear-lanceolate, 
0.5-0.8(-l .6) mm long, 0.6-0.8 mm wide, apically acute to acuminate to 
attenuate, medially thickened, densely and prominently red and black punctate 
medially, with a few scattered rufous papillae, the margin scarious, highly 
erose, densely glandular-ciliate; corolla subrotate, 2-2.3 mm long, the tube 
0.6-0.8 mm long, the lobes suborbicular, unequally divided, 1 .4-1.6 mm 
long, 1.3-1.7 mm wide, apically broadly rounded, densely and prominently 
orange punctate without, densely glandular-granulose throughout within, 
the margin often revolute at maturity, irregular, entire, glandular-granulose; 
stamens ca. 1 mm long, the filaments developmentally fused to the corolla 
tube for their entire length (the stamens appearing epipetalous), 0.6-0.8 
mm long, the anthers very widely ovate, 0.4-0.5 mm long and wide, the 
apically acute, basally cordate, apically dehiscent by termmal pores, confluent 
at anthesis, ventrally sparsely rufous papillate basally, dorsally densely ru- 
fous papillate and sparsely but prominently orange punctate; pistillode absent 
or highly reduced, ca. 0.5 mm long. Pistillate inflorescence as in staminate 
but 3-10(-l4) cm long; peduncle 6-10 mm long; pedicels 1.5-4 mm long. 
Pistillate flowers as m staminate bur 0.7-0.9 mm long, the tube ca. 0. 1 mm 
long, the lobes oblate, 0.6-0.8 mm long, 1-1.2 mm wide; corolla as in staminate 
but 1.4-1.8 mm long, the tube ca. 0.5 mm long, the lobes ovate to subor- 
bicular, 1.1-1.7 mm long, 1.0-1.2 mm wide, the staminodes resembling 
stamens but with antherodes 0.3-0.4 mm long and wide; pistil ellipsoid, 
1-1 .2 mm long, 0.6-0.8 mm diam., the stigma capitate, 3-4-lobed, densely 
translucent glandular-lepidote, ovules 2-3, immersed in the placenta. Fruit 
globose, 5-7 mm long and diam. at maturity, densely and prominently punctate, 
with a few persistent translucent Icpidote scales. 

Distribution. — As here recognized, Cybianthus fnsciis occurs from the Guianas, 
to Venezuela, Colombia, Ecuador, Peru, Bolivia, and their corresponding 
frontiers with Brazil. Cybianthus fliisciis rims the Amazon Basin, from 100- 
200 m elevation. As stated earlier, this is an infrequent distribution, as in 
Cybianthus Venezuela nus. 

Ecology and conservation status. — Cybianthus fuscus occtirs in primary forest 
on terra firnie, especially on steep slopes, near water courses. It is a locally 
infrequent element of the understory but does not appear to be threatened 
at this time. 

Etymology. — The epithet refers to the color of the tomentum of the branchlets, 
pedicels and calyx. 

PiPOLY, Cybianthus in Ecuador and Peru 151 

Represenrarive specimens examined. ECUADOR. Napo: Small area of non-inundated 
forest, ca. 60 km upriver from Nuevo Rocafuerte, 1 3 Sep 1977 (fr), R. Foster 361H (F, USM). 
PERU. Loreto: Prov. Alto Amazonas, Capahuari Sur (Petroleum Camp), 02° 51' S, 76° 
20' W, 200 m, 25 Mar 1982 (fr), R. Va'squez et at. 3065 (AMAZ, MO, US); Prov. Maynas, 
Rio Yavari, Petropolis, 3 km from Rio Amazonas, 8 Sep 1976 (fr),/. ReviIL/ 1302 (AMAZ, 
BRIT, MO); 15 km from roadside along Rd. between UNAP Agricultural Experiment area 
and Escuela Forescal Vivero, 9 Feb 1968 (pist. fl), D. Simpson &J. Schutike 647 (F, USM); 
Puerto Almendras, 03° 45' S, 73° 25' W, 122 m, 7 Dec 1982 (pist. fl), R. Vasquez & N. 
Jaramillo 35! I (AMAZ, MO, NY). 

Cybianthus fusa/s is a widely defined, infrequent, but widely distributed 
species, and is most variable with regard to leaf size and inflorescence stat- 
ure. Populations in Ecuador and Peru are almost identical to specimens known 
from the Guianas, Bolivia, and Brazil in the northeast portion of Amazonas 
state, near the border of Territorio do Roraima. Populations matching the 
type have leaves much smaller than the Ecuadorean and Peruvian popula- 
tions do, and are more like those of the SE Amazon Basin. While Cuiaba is 
located at the northern extreme of the Pantanal Region, it is not entirely 
clear where the exact type locality was. If the type locality in what was Cuiaba 
Province, was north of the Chapada dos Parecis, then it would be at the 
headwaters of the Rio Juruena or Rio Teles Pires, both of which dump into 
the Rio Tapajos, then to the Rio Amazonas. If the locality was west of Cuiaba, 
toward the Bolivian border, streams there form part of the headwaters of 
the Rio Mamase, a branch of the Rio Madeiras, which empties into the Amazonas 
near Manaus. In either case, the type locality would be at the vety edge of 
the Amazon Basin sensu strkto and it would not be surprising to see the species 
in other parts ot the Basin. Therefore, while populations from the type lo- 
cality are slightly smaller in stature, the leaves are more chartaceous, and 
the inflorescences shorter, there is good evidence to show that they are part 
of a large polymorphic ochlospecies complex, of which the populations in 
Ecuador and Peru represent a commonly encountered morphotype. The same 
pattern of variation seen in this species is seen in many Piperaceae (R. Callejas, 
pers. comm.). 

Cybianthus fuscus appears to be closely related to C. a/neifolius Mart, (in- 
cluding C. indecorus Mez), a vicariant species from SE Brazil. The unique 
indument, pedicels obconic in fruit, and striking leaves with very long and 
attenuate apices and bases allow for easy recognition oi Cybianthus fuscus. 

42. Cybianthus cyclopetalus Mez, Bull. Herb. Boissier, Ser. 2, 5:5.33- 1905. 
Type: BRAZIL. Ama/.onas: near Jurua, Miry, Sep 1903 (stam. fl), E. Vie 5840 (uo- 
lotype: B -destr.; leciotype, here designated: FFBG; isolectotypes: G, K, MG). 

Shrub to 1.5 m tall. Branchlets terete, 1.5-2.5 mm diam., densely rufous 
stellate tomentulose, the tomentum appressed. Lmves loosely pseudoverticillate; 
blades chartaceous, elliptic to oblanceolate, (9.5-)l 2.5-l6(-20.5) cm long. 

'^- SlDA 18(1) 

3-5-5(-7) cm wide, apiailly subacuminare to acuminate, basally cuneate, 
decurrcnt thtoughout petiole length, midtib depressed above, prominently 
raised below, decurrent to base of petiole, the secondary veins 10-15 pairs, 
dull and glabrous above, dull and sparsely rufous puberulent below, con- 
centrated along the midrib and the secondary veins, prominently peipuncticulose 
and black punctate-lineate, the margin slightly revolute upon drying, ir- 
regular, entire; petioles marginate and canaliculate, (1-) 1 .5-2(-3) cm long, 
tapered, sparsely stellate rufous puberulent, glabrescent, Staiiiniate infhmcence 
an erect, simple raceme, {4-)9-l 3 cm long, sparsely rufous stellate puberulent; 
peduncle (().6-)().8-l (-1 .3) cm long; floral bracts coriaceous, linear-lanceolate, 
1.3-1.3 mm long, 0.2-0.3 mm wide, apically attentuate to a rounded tip, 
glabrous above, densely and minutely rufous stellate tomentulose below, 
the margin glabrous, entire; pedicels cylindrical, 3.7-5 mm long, sparsely 
rufous stellate tomentulose, glabrescent. Stumhhite flowers 4-merous, coria- 
ceous, nodding, green; calyx 0.9-1. 1 mm long, the tube 0.2-0.3 mm long, 
the lobes linear-lanceolate, 0.7-0.8 mm long, 0.2-0.3 mm wide basally, 
apically long-attenuate, densely and prominently red and black punctate, 
sparsely rufous stellate puberulent, glabrescent, the margin irregular, erose, 
minutely ciliolate; corolla subrotate, 1 .5-1 .8 mm long, the tube 0.6-0.7 
mm long, the lobes very widely ovate, 0.9-1 . 1 mm long, 1 .3-1 .7 mm wide, 
apically emarginate, densely and prominently red and black punctate, gla- 
brous without, densely glandular-granulose througout within, the margin 
irregular, entire, flat, densely glandular-granulose; staminal tube inconspicuous, 
adnate to corolla throughout, 0.6-0.7 mm long, the filaments short, thick, 
0.1-0.2 mm long, glabrous, the anthers widely ovate, 0.5-0.6 mm long, 
0.4-0.5 mm wide, apically and basally trunctae, the connective epunctate, 
glabrous; pistillode subglobose, 0.3-0.4 mm long, 0.3-0.4 mm diam., hollow, 
densely translucent glandular-lepidote, the stigma obsolete. Pistillate inflorescence 
as in staminate but 1-6.5 cm long; peduncle 1-1.5 cm long; floral bracts 
0.9-1 . 1 mm long, 0. 1-0.2 mm wide; pedicels 1.5-2.5 mm long. Pistillate 
flowers as in staminate but calyx 0.8-1 mm long, the tube 0.1-0.2 mm long, 
the lobes, 0.7-0.8 mm long, 0. 1-0.2 mm wide basally; corolla, staminodes 
and pistil unknown. Fr///V globose, 0.3-0.5 mm long and wide, green, exo- 
carp thin, red punctate. 

Distribution. — Cybianthiis cyclopetalns is restricted to the western Amazon 
Basin of Brazil and southeastern Peru, to 290 m elevation. 

Ecology and conservation status. — This species is retricted to varzea or "tahuampa" 
habitats, subject to inundation. Rapid development along the rivers in the 
Amazon Basin changes its flow and may effect these populations. There- 
fore, it should be considered threatened. 

Etymology. — The specific epithet refers to the very widely ovate petals of 
the species. 

PiPOLY, Cybianchus in Ecuador and Peru 153 

Represenracive specimens examined. PERU. Madre de Dios: Prov. Tambopara, Tambopaca 
Wildlife Reserve, 30 km S of Puerto Maldonado, 12° 15' S, 69° 17' W, 260 m, 9 Nov 
1984 (stam. fl), H. Yoiiug etal. 146 (MO, US); Tambopata Reserve, at mouth of Ri'o Orbigny, 
12° 50' S, 69° 17' W, 250 m, 6 Mar 1981 (fr), A. Gentry & K. Young 32028 (MO, USM); 
Along trail Irom large laguna at end of Swamp Trail, Explorer's Inn, near confluence of Rfo 
Tambopata and Rio La Torre, Reserva Tambopata, 12° 50' S, 69° 20' W, 39 km SW of 
Puerto Maldonado, 14 Oct 1985 (stam. fl), S. Smith et al. 738 (US), (stam. fl), D. Bell 101 
(US); Explorer's Inn, Permanent Plots, Tambopata Reserve, 12° 50' S, 69° 17' W, 290 m, 
1 8 Sep I 994 (stam. fl), R. Vchqiiez et cl. 19132 (AiMAZ, BRIT, CUZ, MO) . 

Cybianthus cyclopetalus is most closely related to C. restnosus Mez, but can 
immediately be separated from it by the thinner branchlets, dull, chartaceous 
leaves, and the unique linear-lanceolate calyx lobes with long attenuate apices. 

43.CybianthuspenduliflorusMart.,Nov. Gen. Sp. PI. 3:87. 1831 [1829}. 
Cylmitithm pencliilniiis A. DC, Trans. Linn. Soc. London, Bot. 17:104. 1834 [orth. 
var.}. Cyhicinthin pendiflorin A. DC, Prodr. 8:1 17. 1844 [orth. var.]. Tyfi;: BRAZIL. 
Amazonas: Prov. Rio Negro, near Ega, 170 m, without date, (stam. fl), C. Marti//.f 
s.n. (uolotype: M). 

Cyhicinthiis riiacrophyllus Miq. in Mart., El. Bras. 10:292. 1856. PI. 36. syn. nov. Peckla 
nuicrophylla (Miq. in Mart.) Kuntze, Revis. Gen. PI. 2:402. 1891. T^Ti:: BRAZIL. Amazonas: 
Near Ega, without date, (pist. fl), E. Poepprg 2709 (i.uctotype, here designated: W). 

Shrub or small tree to 4 m tall. Branchlets terete, 2.5-3.5 mm diam., stel- 
late rufuous tomentose, glabrescent. Leaves alternate; blades chartaceous, 
elliptic to oblanceolate, (8-)l().5-19(-28.4) cm long, apically acute, ba- 
sally broadly acute, slightly decurrent on the petiole, midrib raised above 
and below, the secondary veins (10-)12-15(-23) pairs, nitid above, pallid 
below, glabrous, densely black punctate, the margin flat, entire or bearing 
a few rough serrulations; petioles semiterete, (0.5-)0.8-1.2(-1.5) cm long, 
tapered, glabrous. Staminte inflorescence a lax, simple raceme (8-)l()-l4(-2()) 
cm long, densely rufous puberulent; peduncle (().5-)l-1.2(-1.5) cm long; 
floral bracts membranaceous, linear-lanceolate, 1-1.2 mm long, 0. 1-0.2 mm 
wide, apically attenuate, densely rufous puberulent, the margin entire; pedicels 
cylindrical, l-2(-2.5) mm long, densely rufous puberulent, glabrescent. 
S laminate flowers 4-merous, erect, membranaceous green; calyx cotyliform, 
0.8-1.1 mm long, the tube 0.1-0.3 mm long, the lobes ovate, 0.4-0.6 mm 
long, 0.6-0.9 mm wide, apically obtuse, densely and prominently black 
punctate, the margin crenulate, long glandular-ciliate; corolla subrotate, 
1.2-1.5 mm long, the tube 0.3-0.4 mm long, the lobes very widely ovate, 
0.9-1.2 mm long, 1.2-1.5 mm wide, apically obtuse to rounded, densely 
and prominently black punctate, glabrous without, densely glandular-granulose 
throughout within, the margin involute, densely glandular-granulose, en- 
tire; stamens 0.7-0.9 mm long, the tube completely adnate to corolla tube, 
0.3-0.4 mm long, the anthers ovate, 0.4-0.5 mm long, 0.3-0.4 mm wide, 
apically acute to obtuse, basally cordate, the connective prominently black 

154 SiDA 18(1) 

punctate clorsally; pistillode obsolete. Pistillate in florescence as in staminate 
but (2.5-)4-8(-ll) cm long; peduncle (().3-)().5-l cm long; fioral bracts 
1-1.2 mm long, 0.2-0.3 mm wide; pedicels 0.2-0.5 mm long. PistilUite 
flowers as in stammate but calyx 1.2-1.5 mm long, the tube 0.3-0.4 mm 
long, the lobes widely ovate, 0.9-1.2 mm long, L.2-1 .5 mm wide, apically 
rounded, corolla and staminodes unknown; pistil conical, 1 .2-1 .3 mm long, 
0.9-1.1 mm wide, the stigma 4-lobed, the placenta cotyliform, ovules 4, 
naked. Fr/fit 6-10 mm long and in diam., the exocarp thin, densely black 
punctate. Bisexual inflorescence: as in staminate but 6-15 cm long; peduncle 
0.8-1.2 cm long; floral bracts 1-1.2 mm long, 0.1-0.2 mm wide; pedicels 
1-1.2 mm long. Bisexual (lowers as in staminate but calyx 0.7-1 mm long, 
the tube 0. 1-0.2 mm long, the lobes 0.6-0.9 nnm long, 0.5-0.8 mm wide; 
corolla 1.2-1.5 mm long, the tube 0.3-0.4 mm long, the lobes very widely 
ovate, 0.9-1.2 mm long, 1.2-1 .5 mm wide, the margin involute, densely 
glandular-granulose, entire; stamens 0.7-0.9 mm long, the tube completely 
adnate to corolla tube, 0.3-0.4 mm long, the anthers ovate, 0.3-0.4 mm 
long, 0.3-0.4 mm wide, apically acute to obtuse, basally cordate, the con- 
nective prominently black punctate dorsally; pistil conical, 0.9-1.1 mm 
long, 0.7-0.9 mm diam., the style 4-lobed, the placenta cotyliform, ovules 
3, naked. Bisexual fruit globose, 4-6 mm long and in diam., the exocarp 
thin, pellucid punctate. 

Distribution. — Cyhianthus pendidflorus is known from Brazil, Peru and Bolivia, 
100-200 m. The species is not known from Ecuador, but may be expected 
anywhere the habitat is appropriate within the Ecuadorean Amazon. 

Ecology and conservation status. — Cybmnthuspenduliflorus is endemic to igapo 
habitats, and withstands flooding. As a small shrub, it grows on riverbanks 
and on hummocks, behind Triplans (Polygonaceae) and other shoreline plants. 
At this time, it is not considered threatened. 

Etymology. — The specific epithet refers to the lax habit ot the inflorescence, 
pendent in the field. 

RL-pre.sentativc specimens cxiimiiit'd. PERU. I.oreto: Prov. Maynas, Quistococha, 100 
m, 1 Feb 1979 (fr), F- Maid 1623 (AMAZ, MO), Quisrococha, 00° 45' S, 73° IV W, 122 
m, 27 Alii,' 19«7 (scam. H), R. Vauj/zez & N. Jaramillo 9461 (AMAZ, MO, USM); Lower 
Rio Momon, tributary of RTo Nanay, near Iquitos, 8 Dec 1979 (bud), A.Juius & C. Ddvidsim 
9717 (AMAZ, CAS, MO), Near Momoncillo, 1 6 Nov 1976 (stam. i\)J. Rev/t/a 1826 (AMAZ, 
F, MO); Dtto. Iquitos, caseri'o near Nina Rumi, on Rfo Nanay, 23 Feb 1976 (fr),J, Rei'illa 
1S7 (AMAZ, F, MO, USM); Vicinity of Iquitos, 10 Sep- 12 Oct 1976 (bud),/ Revii/a 
1442 (AMAZ, BRIT, MO, USM); Morona Cocha, near Iquitos, 100 m, l4 Dec 1962 (fr), 
_/, Sdwuk' 6268 (AMAZ, F, M(3), UC:LA, US, USM); Puerto Almendras, 03" 48' S, 73° 2V 
W, 122 m, 17 Aug 1983 (bud), /^. Vasquez & N. Jc/nm/tto 4285 (AMAZ, MO, NY, USM); 
Naiica, Quebrada Saragoza, 04° 29' S, 73° 35' W, 1 50 m, 10 Jan 1 988 (fr), R. Va'u/urz & N. 
Janimillo 10339 (AMAZ, MO, US, USM); Iquitos and vicmity, 1 I Oct 1929 (bud), U. 
Willhiim 3(-)76 (F). BOLIVIA. Santa Cruz: Velasco Prov., Campamcnto El Refugio, along 
Rio Paragtia, SF of the house, M° 46' 09" S, 61° 02' 1 I " W, 240 m, 1 1 Oct 1994 (H bud), 

PiPOLY, Cybianrhus in Ecuador and Peru 155 

R. Giiillm & G. Scilvatierni 2290 (BRIT, MO, USZ); Campamento La Toledo, 1 ,()()() m E of 
the house, 14° 42' S, 61 ° 09' W, I6()m,21 Oct 1994(sram. fl), /?, G//!!le>i&R. Chore2439 
(BRIT, MO, USZ), 1 km W of camp, on canoe route to Campamento Toledo, 14° 45' 51" 
S, 61° 02' 22" W, 30 Jan 1995 (fr), R. Guillen et al. 3114 (BRIT, MO, USZ). 


Support for my studies on the Flora of Peru was provided by a generous 
grant from the Andrew W. Mellon Foundation (during my tenure at the 
Missouri Botanical Garden), which is gratefully acknowleged. I would also 
like to thank the John D. and Catherine T. MacArthur Foundation for their 
support of fieldwork in Peru as part of the Documentation of Neotropical 
Plant Diversity Project, which I directed from 1990-1994. Thanks are also 
due to Michael Dillon (F) and the late Al Gentry (MO) for their collabora- 
tion with the Flora of Peru project, to Enrique Forero (COL) and Warren D. 
Stevens (MO) whose advice was invaluable. Many thanks to Jon Ricketson 
(MO), then project coordinator and now colleague in Myrsinaceae system- 
atics and authority on the systematics of Parathesis. For their warm hospi- 
tality and collaboration, whether in the field in the herbarium, I express 
my sincere gratitude to Rodolfo Vasquez (MO), Camilo Dfaz (MO), Nestor 
Jaramillo (MO), Cesar Grandez (UNAP), Rosa Ortiz de Gentry (MO), Martin 
Timana (TEX), Abundio Sagastegui (HUT), Jose Mostacero (HUT), Joaquina 
Alban (USM). My volunteer at MO, Catherine Mayo, assisted in the move- 
ment of specimens and recording of determinations. Studies on Cyhianthus 
subgenera Microconomorpha. Conomorpha. Triadophora and Cornomyrsine were 
completed during my tenure as a Postdoctoral Fellow in the Department 
of Botany, National Museum of Natural History, Smithsonian Institution. 
My continued status as a Smithsonian Research Associate has permitted me 
to continue studies in the group, particularly in their strong Peruvian and 
Colombian collections. I thank Linda Ellis, Peggy Duke, Juan Pinzon, and 
Linny Heagy for the skillfull illustrations of the new taxa described herein. 
I am most grateful to the directors and curators of herbaria cited, especially 
F, G, GB, K, MO, NY, S, and US, for their patience and work in arranging 
for loans and various transfers of material. Finally, to Jon Ricketson (MO), 
and Ted Barkley (KSC) for their painstaking review of the manuscript and 
helpful comments, and to Barney Lipscomb for his unselfish, time-consuming, 
and meticulous work in editing, I owe my sincerest gratitude. 


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Ph. D. Thesis. The City University of New York, New York. 

1980. Una nueva clasificacion del genero Cyhianthus (Myrsinaceae). Acta 

Biol. Venez. 10:129-185. 

156 SiDA 17(4) 

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Gross, J. 1972a. Die Uiltblatter der Nymphaeaceen. Ber. Detitsch. Boc. Ges. 45:454-458. 

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1. Cybianthus pastensis 24. 

2. C. t^igantophyllus 25. 

3. C. occigranatensis 26. 

4. C. spichigeri 27. 

5. C. lepidotLis 28. 

6. C. laetus 29. 

7. C. peruvianas 50. 

8. C. tomperuvianus 30a. 

9. C. guyanensis subsp. pseudoicacoreus 30b. 

10. C. cimanae 31. 

11. C. CLiarrecasasii 32. 
1 2. C. nestorii 33. 
1 3. C. spicatus 34. 

14. C tulvopulverulentus subsp. magnoliifolius 35. 

15. C. verticilloides 36. 
I 6. C. LToarii 37. 

17. C. humilis 38. 

18. C. sprucei 39- 

19. C. simplex 40. 

20. C. kayapii ■• 41. 

21. C. anrhLiriophyllus 42. 

22. C, schlimii 43. 

23. C. poeppigii 

C. pseudolongitolius 
C. vasquczii 
C. cenepensis 
C. nanayensis 
C. marginatus 
C. lineatus 
C. magnus 

C. magnus subsp. magnus 
C. magnus subsp. asymmetricus 
C incognitos 
C. mmutifiorus 
C. huampamiensis 
C. granulosus 
C. fjavovirens 
C. vcnezuelanus 
C. grandezii 
C. jensonii 
C. foscen 
C. rcsinosus 
C. tuscus 
C. cyclopetalus 
C. penduliflorus 

158 SiDA 18(1) 


Figures in parentheses refer to numbers from the numerical Hst of taxa. 
Collecrion numbers in boldface type indicate type specimens. 

Acnsca-Solis, M. 5544 (20); At;Lnlar, M^ & D. Casrro 623 (7); 655 (7); ,S()5 (9); Albert clc Escobar, L. 
L't al. 3744 (16); Alvarez, A. L-t al. 381 ( 1 ); 4 1 2 ( I ); 490 (1); 1 330 ( 1 ); Anciiash, E. 93 (8); Itl (8); 
21 1 (20); 220 (S); 274 (33); 392 (33); 522 (26); 580 (2); 588 (33); 731 (33); 1405 (20); 1412 (26); 
1437 (2); Antlrc, E. s.n. (22); 1151 (22): 3819 (17); 45 51 (1); Arias, L. et al. 134 (22); Arroyo, L. & 
K, Kciil l6-i (S); Arroyo. L. ct al. 510 (9); 67 'i (8); Asplund E. 9396 (20); 10209 (2i); Aspluiid, E. 
10302 (23); 12197 (23); 18717 (2])\ Aulesna, C. & M. 1313 (3); Aulcstia, M. & J. Andi 925 (9); 
Ayala, 1-. 1623 (43): 2102 (36); Ayala, I", cc al. 2814 ( t6). 

Balsk-v, H. 4295 (I); Balslev, 11. & -Santos Dt-a 2850 (15); Bang, M. 2048 (36); Barbour, P. 2405 
(31); Barbour, P. 2567 (31); Bell, D. 101 (42); Berlin, B. i93 (20); 1760(33); 779(33); Betancur.J. 
et al.4857 (19); Bohlin, J. et al. 1493 (18); Boyle, B. 3 173 (19); 3599 (19); Boyle, B. &J. Brailtord 
1878 (I 9); Boyle, B. et al. 337 i (28); 3450 (30b); Bradford, J. et al. 55 (1 ); Brandbyge, J. 42095 (1); 
Brandby.ue, J. & Barlord, A. 42506 ( 1 ); Bravo, E. & R Gdmez 19 (2 I ); Brenes, A. 20530 (22); Buclitein, 
O. 175.^ (8); 1758 (5); Bunting, G. & I.. Licht 775 (22): Burger, W. & R. Stoize 5803 (22). 

Calderon, C. et al. 2852 (22); Gillejas, R. et al. 2737 (3); Camp, W. E-4809 (28); Campos, J. & L. 
3121 (28);Campos,J.etal. 3161 (6); Castaneda, R. 2873 ( IS); 4746 (22); Castro, J. 27 (7);Cer()n,C. 
2494 ( I 8); 2585 (23); 7 1 84 (17); 7409 ( I 8); Cerdn C. & M. Faetos 764 1 (20); 76>48 (18); Cerdn, C. & 
N. Gallo 5063(9);Chat(')n, I. &(i. Herrera I 720 (22); Cogollo, A. & C. Estrada 296 (3); Cogollo, A. 
etal. 7529(3);Collenete, B. 1 1 3 (8); Cornejo, X. & C. Bonifaz 2979 (19); 4339 (19); Croat, T. 7115 
(18); 17 167 (22); 19485 (38); 20528 (U); 21 194 (8); ^7785 (23); 49361 (20); 59009 (16); 72394 
(19); 73352 (I H); Croat, T. & A. Cogollo 52257 (18); Cuatrecasas, J. 14053 (18); 14138(18); 14918 (3); 
14947 (30a); 15474 {22}\ 15658 (3); 15743 (18); 16272 (20); 16272 (18); 17312 (22); 17625 (18); 
19892 (18); 21721 (15); 21805 (28); 21981 (15); 22131 (3); 23583 (I I); 23734 (3); 23855 122). 

Daly, D. etal. 5926(1 1 ); de Nevers, (i. & H. Herrera 6642 (22); Di'az, C. 566 (22); 7265 (20); 1474 
(31); l486(4());Dfaz,<;:. &J. Campos 37 11 (l);Di'az, C. & A. Torres 7805 (30a); Dfaz, C. etal. 3330 
(1); 4448 (1); 6930 (3.^); 6993 (7); 7007 (2); 71 70 D; ^252 (5); 7585 (30b); 7649A (34); Dillon, M, 
eral. 1 2 1 9 (8); Dodge, C:, & V. Cioerger 9283 (22); Dodson, C. 2821 (7); Dodson, C. er al. 7597 (19); 
9101 (19); 151 15 (3); Domfnguez, F. 147 (20); Dressier, R. 6036(20); Drew, W. E-351 (28); Dudley, 
T. 10668 (30b); 10690 (6); 10803 (6); 10910 (28); 10922 (28); 10931B (28); 11324 (30b); 1 1915 
(1); 13124(13); 13513 (1). 

Encarnacitin, R 864 (7); 26200 (1); Espinal, S. 2109 (22). 

Faber-Eangendoen, D. & J. Htirrado 1757 (40); I'erreyra, R. 1()40 (32); E'lgueroa, A. 875 ( I 5); b'oLsom, 
J. 4544 (18); Fonnegra, R. et al. 3076 (2i); I'orero, E. & L. Wriglcy 7035 (8); 1325 (22); Forero, E. 
et al. 7350 (22); Eosberg, F R. 1 9969 (ID; 2()li8 (22); 22376 (II); Fosberg, R R. & Giler, M. 23 1 19 
(I); Foster, R. i618 (W); 3708 (26); 4226 (4); 4243 (27); 4327 (27); 4404 (40); 7817 (32); 8624 
(32); 898 1 (32); 9013 (30b); Foster, R. & T'. Watcher 7242 (39); 7254 (39); Foster, R. & B. d'Achille 
10013 (23); Foster, R. 8962 (8); 10860 (1); 10993 (9); Franco, R et al. 1059 (22); Franco, R et al. 
3204 (17), 15423 (23); Funck, N. & L. Schlim 678 (36). 

Ciarci'a, I: &J, Echavarna 259- A (20); Garcia, E &J. Ecliavarria 259- A ( 1 7); Gehriger, W. 471 (28); 
Gentry, A. 124 19 (30b); 20763 (27); 80096 (4); Cientry, A. & J. Aronson 25044 (2); Gentry, A. & L. 
E.mmons i8776 (27); Gentry, A. & E. Forero 73 17 (18); Gentry, A. & N. Jaramillo 22 1 3 1 (40); Gen- 
try, A. & A.Juncosa4l702 (22); Gentry, A. & S.Mori 13665 (23); 13709(23); 1 4023 (23); Genrr> 
A. & G. Shupp 26565 (19); Gentry, A. & J. Solomon 44407 (5); 44668 (5); C3entry, A. & K. Voting 
32025 (27); 2028 (42); Gentry, A. ct al. 16842 (3); 16867 (3); 21033 (27); 22367 (27); 22859 (33) 
22859 (tI); 2291 1 (31); 27952 (2); 29700 (38); 31479 (27); t5062 (20); 35188 (19); 39040 (7) 
40453 (18); -il88() i2i)\ 42183 (9); ■I22i7 (20); 51083 (36); 51400 (36); 53167 (3); 53326 (22) 
53717" (18); 5>S81 (11); 5 3960 (11); 5 5053 (19); 55708 (38); 72129 (9); 72499 (19); 74630 (10) 
80179 (4); 56531 (4); 53551 (3); Gleason, H. 159 (36); Grandez, C. et al. 1824 (37); 4487 (9) 
4711 (7); 5321 (12); 5370(12); 5642 (12); Gravum. M. & G. Sharz ^170(22); Grubb, R etal. 744 

PiPOLY, Cybianthus in Ecuador and Peru 159 

(6); 1073 (1); 1633 (20); Guidino, E. 137 (7); C.uillcn, R. & G. Salvatierra 2290 (43); Guillen, R. & 
R. Chore 2459 (43); Guillen, R. et al. 31 14 (4 3); Gutierrez, G. 1 186 (23). 

Haenke, T. 98 (1); Hammel, B. 17150 (19); Harling, G. 25313 (1); 25334 (1); Harling, G. & L. 
Andersson 1725 5 (23); 18778 (19); 21594 (1); 23540(1); 23842 (1); 24097 (23); 9764 (23); Harcman, 
R. 2401 (23); 12461 (23); Hartshorn, G. 2996 (35); Hartweg, C. 1200 (28); Haught, O. 1502(22); 
4660 (22); 5479 (22); Hernandez, J. & S. Hoyos 483 (22); Holm-Nielsen, L. 681 8 (1); Holm-Nielsen, 
L. eraL 3965 ( 1 ); 29949 (28); 19295 (1 8); Hoover, W. 1 194 (22); Hoover, W. et al. 2456 (1 9); 2809 
(19); 2815 (22); 3358 (19); Huashikat, V. 356(20); 507 (20); 581 (20); 677 (34); 1221 (34); 1248 
(23); 1422 (34); Humboldt, A. von & A. Bonpland 1096 (13); Hurtado, K & D. Neill 235 (3); 
Hurtado, F. & A. Alvarado 503 (20). 

Idrobo, J. et al. 9671 (17); Imthurn, E. B/9 (14). 

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160 Si DA 18(1) 

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Botanical Research Institute of Texas 

do Papua New Guinea Forest Research Institute 

R 0. Box 314. Lae. 4lh PAPUA NEW GUINEA 


Botanical Research Institute of Texas 

309 Pecan Street 

Fort Worth. TX 76102-4060 U.S.A. 

jpipoly @ brit. org: clusia@latinmail. com 


Botanical exploration of the Hans Meyer Range in southern New Ireland revealed the 
existence of two previously unknown taxa in the families Rubiaceae and Corsiaceae. The 
novelties, Psychotria osiana Takeuchi & Pipoly, (Rubiaceae) and Corsia piirpurata var. iviakabui 
Takeuchi & Pipoly (Corsiaceae) are described and illustrated. Taxonomic and ecological 
notes are also provided for the new taxa. 


Wanpela wok bus long salt bilong botani ibin kamap long ol lain maunten bilong Hans 
Meyer, Niu Hand Province, long yia 1994. Dispela wokbus o stadi ibin kamapim tupela 
niupela samting. Wanpela em sotpela diwai bilong famili Rubiaisi. Namba tu em wanpela 
liklik gras nating bilong famili Korsiasi. Tupela igat nem olsem: Psychotria osiana na Corsia 
piirpurata var. wiakabui. Dispela stori i tok klia na soim tu sampela piksa bilong tupela. 


Papua New Guinea (PNG) is one of only four countries on earth projected 
as retaining most of its original forest cover by the end of this century (Suzuki 
1993). Unfortunately, it is also among the least known floristic areas within 
the Malesian region (Conn 1994; Johns 1995). In the recent multiagency 
Conservation Needs Assessment (CNA), 17 localities of primary conserva- 
tion significance were identified for Papuasian forest environments (Beehler 
1993). Southern New Ireland was one of the enumerated sites considered 
vital to biodiversity preservation and in critical need of current informa- 
tion. As a direct consequence of the CNA evaluation, a biological assess- 
ment expedition was organized by Conservation International and the Papua 
New Guinea Department of Environment and Conservation (DEC) in January- 
February 1994. A multidisciplinary compilation of results from that sur- 
vey has been prepared as a Rapid Assessment Protocol (RAP) Working Paper 

SiDA 18(1): 161-168. 1998 

162 SiDA 18(1) 

by Conservation International (Beehler, in press). Orchid specimens from the 
Hans Meyer trip were previously reviewed in a separate speciaUst account 
(Howcroft 1994). Two new taxa discerned during general examination of 
the expedition's botanical gatherings are described here. 


Psychotria osiana Takeuchi Sc Pipoly, sp. nov. (Fig. 1). Typf; PAPUA NEW 
(iUINHA. NiAV Ireland: Hans Meyer Range, pond next to 'Lake Camp,' ()4° 27.205' 
S, 152" 56.489' E, 1,175 m, 29 Jan 1994 (tl, fr), W. Takei/chi &J. Wiakabu 9650 
(iiOLOTVPF: LAE; isotyphs: A, BRIT, K, L). 

Quoad scipLila elongata calyptrataque, stipula ab laminis juveniliLis rumpentes, P. leLiiinides 
valde arete affinis, sed ab ea inttorescentiis trichotomiis reduciseiue (non monoromiLs eiongacLsque), 
floribus verticellatis (non alternatis) praeditis, lamini.s secLis costis iurluraceo-lepidotis (non 
glabris) denique petalis desuper papiUosis (non glabris) statim cognoscitur. 

Shrub or small tree to 7 m height, 8 cm dbh. Stem straight, isodiametric, 
basal swell absent; outer bark brown, smooth, slash and sapwood stramineous 
to pale yellow. Branchlets terete, 2-3 mm diam., pale brown, slightly com- 
pressed at the summit, laxly pilose at apical nodes otherwise glabrescent. 
Stipules sheathing, calyptrate, caducous, translucent whitish-green, glabrous, 
to 5.5 cm long and 6 mm wide. Leaves opposite, usually conferred; blades 
coriaceous, elliptic, 10-13 cm long, 3—4 cm wide, apically acuminate, ba- 
sally attenuate, the margin entire, adaxially nitid and glabrous, abaxially 
light green and furfuraceous-tomentose along the midrib, the midrib prominulous 
on borh sides, more elevated beneath; the secondary veins 10-13 oblique 
lateral pairs, excurrently arcuate and usually closing submarginally, the tertiary 
reticulum lax, irregular, bifacially raised; petioles adaxially plane, 10-20 
mm long, glabrous. Inflorescence a sessile, terminal, pinnate panicle, ruptur- 
ing through rhe stipule, trichotomous (with three principal branches at base), 
the branches with flowers racemose, the primary rachis to 2 cm long, greenish, 
somewhat angulate, laxly pilose at nodes, internodes subglabrous or pu- 
berulent; floral bracts inconspicuous, not persisting; pedicels 1—3 rnm long. 
Flowers opposed or in verricels, 5-merous, entirely glabrous, apparently iso- 
morphic; calyx broadly cupuliform, .5 mm long, 1—2 mm wide, limb den- 
ticulate; corolla tubular, 3 nim long and obtusely cylindrical in bud, the 
lobes chartaceous, corniculate, elliptic to ovate, 1 .5 mm long, 1 mm wide, 
spreading or reflexing at anthesis, adaxially papillose; stamens erect, epipetalous; 
filaments 1 .5 mm long, adnate to rhe corolline sinuses or slightly below, 
the anthers basifixed, oblongoid, 0.7 mm long, 0.2 mm wide, apically and 
basally emarginate; ovary umbonate, channelled, bilocular, the style bifid, 
slightly exserted, the stigmatic lobes spreading and tuberculate. Drupelet 
subglobose, 5—8 mm diam., epicarp glabrous; pyrenes 2, not dorsally ridged, 
approximately planoconvex, flat on the commissural face, often with one 
pyrene aborted or reduced; endosperm ruminate. 

Takf.uchi and Pipoly, New taxa from Papua New Guinea 


Fig. 1 . Psychotria osiana Takeuchi & Pipoly. A.Branchlet. B. Developing inflorescence emerging 
through stipule. C. Inllorescence; showing main rachis and parr of the second rachis in 
back. Third axis deleted for clarity. D. Flower at anthesis; petals reflexing. E. Partially dissected 
flower. A— D drawn trom holotype. 

164 Sum 18(1) 

Distribution aiicl ecoloii^y. — Known only from mossy montane forest in the 
Hans Meyer Range of southern New Ireland. Occurring as a serai element 
in gap phase regeneration among mature growth premontane forest stands, 
and along forest margins. 

Etymology. — It is a great pleasure to dedicate this species to Dr. Osia Gideon, 
Deputy Director of the Papua New Guinea Forest Research Institute and a 
specialist in Papuasian Rubiaceae and Zingiberaceae, for liis relentless ef- 
forts to document the rich, largely autochthonous, flora of New Guinea. 

PakatypI': PAPUA NEW GUINEA. New Ireland: Hans Meyer Range, pond next to 
'Lake Camp,' 04° 27.205' S, 152° 56.489' E, 1,175 m, 27 Jan 1994 (fl, fr), W. I'akeiichi & 
J. Wuikabu 9560 (A, BRIT, K, L, LAE). 

Papuasian Psychotria were first revised by Valeton ( 1 927) and the nonclimbing 
representatives more recently treated by Sohmer (1988). Infrageneric rela- 
tionships are still poorly understood due to the taxonomic difficulty of the 
genus and the absence of regionally-based studies. However a number of 
informal species groups have been recognized by Sohmer from macroscopic 
features such as stipule form and inflorescence structure. 

Psychotria osiarui is characterized by a contracted trichotomous inflores- 
cence (i.e., with 3 rachises branched at the base), flowers opposed or verticelled 
along the main axes, leaf blades with abaxially furfuraceous-tomentose midribs, 
completely glabrous flowers, and petals adaxially papillose. The corolline 
papillae are apparent in fresh or rehydrated material, but less so i>i sicco. 

A total of 17 species of Papuasian Psychotria have sheathing stipules of 
the sort present in Psychotiria osiana. Developing inflorescences and young 
leaves emerge by rupturing through the stipule, which subsequently disin- 
tegrates or falls away intact. The combination of calyptrate (sheathing) stipules 
and a trichotomous inflorescence occurs in only two species: P. leleanoides 
Sohmer and P. lorentzii Valeton. Psychotria osiana is clearly related to P. leleanoides, 
(a species from New Britain and the Solomon Islands), but is distinguish- 
able from it by the leaf indumentum and the entirely glabrous flowers with 
papillate corolla. Judging from elevational distributions, Psychotria osiana 
is the montane sister species to P. leleanoides, the latter being primarily a 
lowland taxon from elevations below 200 m (Sohmer 1988). 


Corsia purpurata L.O. Williams var. wiakabui, Takeuchi & Pipoly, var. 
nov. (Fig. 2). Type: PAPUA NEW GUINEA. New Ireland: Hans Meyer Range, 
pond nexr to 'Lake Camp,' 04° 27.205' S, 152° 56.489' E, 1,175 m, 28Jan 1994 (fl, 
fr), W. Takeuchi &J. Wiakabi/ 96! 1 (holotype: LAE, in spirit). 

Ad Corsia piirpuratam var. purpuratam accedens sed ab ea labello acuminate nee ad basem 
extendens praeclare di.stat. 

Terrestrial herb to 13 cm height, sciophytic, erect, all parts glabrous and 

Takeuciii and PiPOLY, New taxa from Papua New Guinea 


5 mm 

Fig. 2. Carsia puypiiratci L.O. Williams var. iviakabui Takeuchi & Pipoly var. nov. A. I labit, 
dorsal view. B. Habit, lateral view. C. Frontal aspect, petals and lateral sepals. D. Median 
sepal. E. Petal. F. Basal callosity, from side. A-F drawn from holotype. 

166 SiiMlS(l) 

dull reddish-brown. Stenis rerete, weakly sulcate, the nodes distant. Leases 
4-5, spirally arranged, acroscopic, base sheathing; lamina linear-acuminate, 
conduplicate or with margins broadly revolute, chartaceous, 9—17 mm long, 
4—6 mm wide, to 6-nerval, sometimes apiculate; venation visible as dark 
lines, not raised on either side. Flowers solitary, terminal on the unbranched 
stem, bisexual; median sepal flabellate, symmetrical, 16-1 9 mm long, 20- 
23 mm wide, nodding at anthesis, unguiculate, bluntly acuminate at the 
apex; the basal callosity linear, narrowly raised by 4 mm above the plane of 
the 'labelkim,' summit corniculate, base abruptly truncate at the claw; median 
sepal 1 2-veined, the veins paralleiodromous, diverging mainly from the proximal 
half of the sepal, once or several times furcate before the margin; petals and 
lateral sepals isomorphic, basiscopic and incurved, costate, lanceolate, 4 mm 
long, 3 mm wide, acumen 1.5 mm long; stamens 6, in 2 whorls opposite 
the perianth segments; filaments resembling the style, 0.6-0.7 mm long; 
the anther cells 2, oblongoid, 1 .2 mm long; style simple, cylindrical, ca. 1 
mm long. Pr/nt not seen. 

Distribution and ecology. — Known only from the type locality in the Hans 
Meyer Range on New Ireland. Corsia pi/rpunitci var. wicikahiii occurs infre- 
quently in mature-growth, primary premontane forest, on its floor, cov- 
ered by leaf duff. This new variety was encountered as a population flush- 
ing after recent rains and thereafter evanescent; all individuals were seen in 
comparable maturational states. 

Etyiiiology. — The new variety is dedicated to Joseph Wiakabu; botanist, 
explorer-collector, and colleague from the Papua New Guinea National 

Paratvpi:: PAPUA NEW CUIINEA. Ni:\v Huns Meyer Range, slopes above 
river valley on the ascent from Mandih lake, ()4^ 26' S, 1 52^^ 59' E, 7^0 m, 10 Au.^ 1975, 
M.J.S. Scnuh et cil. 2091 (K, EAE). 

Corsia was revised by van Royen { 1 972) in a monograph based on 45 exsiccatae 
numbers, an average of less than 2 collections per species. The limited number 
of specimens available for study is due to population rarity, the inconspicu- 
ous habit of the plants, an ephemeral [4ienology, and the preference for sheltered 
microsites in forest where ambient light conditions favor concealment. Van 
Royen (ibid.) commented that encounters with Corsia are often of a fortti- 
itous nature; occasioned by understory sun flecks falling on the plants in a 
certain way by chance. There is no doubt that the genus is rarely found by 
collectors. The intervening years since the initial revision has seen little increase 
in availability of specimens, nor is the situation likely to improve dramati- 
cally in the future. The difficulty of botanizing purposefully for Corsia dis- 
courages the sort of field-based studies necessary for understanding the 
morphological variation between populations. 

Takeuchi and PiPOLY, New taxa from Papua New Guinea 1 67 

According to van Royen's (1972) conspectus, Corsia is a saprophytic ge- 
nus with 25 species distributed through New Guinea, the Bismarck Ar- 
chipelago, the Solomon Islands, and Australia. Most of the species are nar- 
rowly defined endemics of montane forest habitats, and appear to be restricted 
to specific mountains. On the basis of the venation and basal callosity of 
the median sepal, the new taxon is related to C. purpurata var. purpurata, 
but is distinguishable by the flabellate 'labellum' with acuminate apex and 
the base truncate rather than extended. 

The type locality for var. waikabui is floristically depauperate in com- 
parison to similar habitats from mainland New Guinea, and is dispropor- 
tionately composed of epiphytic or nonendemic species (Takeuchi and Wiakabu, 
in press). Depauperate levels of biological diversity in this montane envi- 
ronment was also reported by zoological specialists participating in the New 
Ireland survey (Beehler in press, passim). The expedition's collective find- 
ings are consistent with a supposition that the cloudy uplands of southern 
New Ireland are of geologically recent origin. In view of the multidisciplinary 
results from the recent survey, it is unlikely that future exploration of the 
Hans Meyer Range will yield significant numbers of additional novelties. 


The Rapid Assessment Protocol survey of New Ireland was funded by 
Conservation International and the Biodiversity Support Program. Expe- 
dition leader Dr. Bruce Beehler and the Papua New Guinea Department of 
Environment and Conservation performed key coordinating roles. The Liz 
Claiborne and Art Ortenberg Foundation and the John D. and Catherine 
T. MacArthur Foundation have provided ongoing financial support to W. 
Takeuchi for studies partially based at the Papua New Guinea National 
Herbarium, at the PNG Forest Research Institute, m Lae, in support of his 
principal work at the Crater Mountain Wildlife Management Area. 

Neville Howcroft prepared the illustration o{ Corsia purpurata var. wiakabui 
and Talk Iwagu the illustration of Psychotria osiana, which we gratefully 


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porate Press, Inc., Landover, Maryland. 

Bi-EHLr,R, B. (ed.). In press. Rapid Assessment Program Working Papers, No. 1 0. A biodiversity 
assessment of southern New Ireland, Paptia New Guinea. Conservatif)n Internatiorial. 
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Conn, B.J. 1994. Documentation of the flora ot New Guinea. In C.-I Peng and C.H. Chou, 
eds. Biodiversity and terrestrial ecosystems. Institute Bot., Acad. Sinica Monogr. 14: 123— 1 56. 

Howc;roft, N.H.S. 1 994. Orchid collections from New Ireland. Orchid Res. Bull 3. Papua 
New Guinea Forest Research Institute. 

168 SiDA 18(1) 

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SoilMF.R, S.H. 19H(S. The nonclimbing species ot the genus Psychntric/ (Rubiaceae) in New 
Guinea anci the Bismarck Archipelago. Bishop Mus. Bull. Bot. 1:1—3.^9. 

Suzuki, l^. 1993. Time for a change. Allen and Unwin, St. Leonards. 

Takeuciii, W. and J. Wiakabu. In press. A transect-based floristic reconnaissance of sourh- 
ern New Ireland. In B. Beehler, ed. A biodiversity assessment of southern New Ireland, 
Papua Nfew Gtnnea. Rapid Assessment Program Working Papers 10, Conservation In- 
ternational, Washmgton, D.G. 

Valhton, T. 1927. Die Rtibiaceae von Papuasien. 11. /.weiter Teil: Cofleoidcae. Bot. Jahrb. 






The University of Western Ontario 

Department of Plant Sciences 

London, Ontario, CANADA N6A 5B7 


Botanical Research Institute of Texas 

309 Pecan Street 

Fort Worth, TX 76102-4060, U.S.A. 


Three new species of Crataegus, C. okennonii J.B. Phipps (from series Douglasianae), C. 
okanagane)isis J.B. Phipps & O'Kennnon (from the newly described series Pi/rpureofri/cti J.B. 
Phipps & O'Kennon), and C. phippsii O'Kennon (near series Molles) are described from western 
North America. All are striking, widespread taxa with a substantial number of records. 

Key Worus: Taxonomy, Crataegus, Rosaceae, new species 


Tres nuevas especies de Crataegus, C. okennonii J.B. Phipps (de la serie Dauglasianae), C. 
okanaganensis J .B . Phipps & O'Kennnon (de la nueva serie descrita Purpureofructi J.B. Phipps 
& O'Kennon), y C. phippsii O'Kennon (proximo a la serie Molles) se describen del oeste de 
Norte America. Todos son taxa sorprendentes y de amplia difusion con un sustancial numero 
de ciras. 

Recent fieldwork with R.J. O'Kennon of Fort Worth, Texas in the American 
states west of the Rocky Mountains and in British Columbia for the first 
author's revision of the red-fruited Crataegi (Phipps 1998) of this region 
has serendipitously uncovered three new species of hawthorn. The first, Crataegus 
okennonii, a new purplish-fruited taxon of ser. Douglasianae (Rehd. ex Sarg.) 
Rehd., striking and obvious in the field, is also usually easily resolved in 
the herbarium. Crataegus okanaganensis, rarely collected before our studies, 
is also a striking new taxon with brilliant red (late August) to deep purple 
(late September) fruit. It is accorded a new series, Pupureofructi, together 
with the long-described and long-overlooked C. williamsii Eggl. The third 
new species, C. phippsii, though first collected as far back as 1908, remained 
otherwise unnoticed until our fieldwork. It is a very distinct purple-fruited 
species related to series Molles. 

SiuA 18(1): 169-191. 1998 

170 SiDA 18(1) 

One of the reasons that these new Crataegus species west of the Rocky 
Mountains have been overlooked has been the fliiltire of nearly all collectors 
to take note of fruit color (Phipps 1998). Each of the following species is 
quite distinct in this respect and we therefore take the opportunity to present 
color plates of the three species illustrating this point. Hawthorns are con- 
sidered to have mostly black or red fruit. The discovery of these three ex- 
tends the number of those which are burgundy to deep purple at some stage 
in their development. Another feature emerging from this work are new 
centers of diversity for western North American Crataegus appearing m the 
northern Okanagan of British Columbia and northwest Montana. 

A further reason that these taxa have been overlooked is that in the Pa- 
cific Northwest region (to nw Montana) and British Columbia where they 
occur there has been a belief that generally, only one red-fruited native spe- 
cies, C. 'Columbiana Howell,' one black-fruited native taxon, C. douglam Lindl. 
(inch C. suksdorfii (Sarg.) Kruschke) and one introduced red-fruited taxon 
(C wrwog}'Wi:7jacq.) occurred. This belief has persisted (Phipps 1998) in spite 
of the widespread presence in this region of C. niacracantha Lodd. ex Loud. 
In view of this lack of taxonomic inattention, including the disappearance 
from the floristic literature of the distinctive C. icillianis'n, it is perhaps not 
surprising that earlier botanists did not recognize potential new taxa in this 
region. Consequently, we take this opportunity of reviewing the main western 
North American species (west of the Rocky Mountains) in a numerical taxonomic 
analysis. This analysis uses one synthetic OTU per taxon, each described 
from typical material over 38 morphological characters (Table 1) of the kind 
commonly used in Crataegus taxonomy. The eleven native species generally 
recognized in the west are all present, together with C. monogyna, a useful 
outgroup, and five species from east of the Rocky Mountains (C. mollis (Torr. 
& A. Gray) Scheele, C. suhmolUs Sarg., C. brachyaeantha Sarg. & Engelm., C. 
schuettei Ashe and C punctata Jacq.) to give structure and scale. Also the 
three new taxa described in this paper are included. A dendrogram was prepared 
by NT-SYS using the options taxonomic distance and unweighted pair-group 
arithmetic averaging. The results (Fig. 1) place C. inonogyiia and C. punctata 
outside all the other species. Among results of general interest, varieties 
(C chrysocarpa Ashe var. chrysocarpa and C. c. var. piperi (Britton) Kruschke) 
cluster at about the 0.6 level while distinct species cluster at the 0.8 or greater 
level, e.g. C. schuettei 'dnd C. wnotoniana Eggl. (series Tenuifoliae), C. suksdorfii 
and C. do//glasii {'iQUi^s Douglasianae) and C. erythropoda Ashe and C. rivularis 
Nutt. (series Cerrones). Other pertinent results are discussed under the new 
taxa described in this paper, 


1. Crataegus okennonii J.B. Phipps, sp. nov. (Fig. 2). 'Ywv. CANADA. 

Piiipps AND O'Kennon, New species of Crataegus from Western North America 171 

TAiii.n 1. The 38 characters scored for tlie numerical caxonomic analysis. 

PLoit. gctiercil 

1 . Plant habit 

2. Bark type 

3. Branch arrangement 

'r barns and I'uigs 

4. Thorns: indeterminate present? 

5. Thorns: length 

6. Thorns: curvatLire 

7. Thorns, color: browns 
(S. Thorns, color: gray 

9. Twigs, I yr old, color; browns 
10. Twigs, I yr old, color; gray 

Lamina: length 

Lamina: length/breadth 

Lamina: position ot widest part 

Lamina: venation, number 

Lamina: veins to sinuses 

Lamina: lobe number 

Lamina: lobe shape 

Lamina: max. sinus depth (LII) 

Lamina: abaxial pubescence, young 

Petiole: glands 

Leaf teeth: glands 














22. Flower: number 

23. Pedicel: pubescence 

24. Anthesis time 


25. Diameter 

26. Calyx lobes: margins 

27. Stamen: no. 

28. Anther: color 

29. Style: numbei 


30. Fruit: color, month before lull ripeness 
3L Pfuit: color at full ripeness 

32. Fruit: shape, 1 

33. Fruit: shape, 2 

34. Fruit: pubescence 

35. Fruit: calyx orientation 

36. Fruit: length (height) 

37. Pyrenes: lateral faces 

Autumnal joltage 

38. Color 

British Columbia: Okanagan Valley, E side of 97A 0.5 mi N of intersection with 
97, scrubby slopes with long cleared area, 50°2rN, 119°15'W, 19 Aug \99A,Phipps, 
J.B. & (rKennon, R.J. 6998 (hoi.otype: UWO; isotypes: BRIT, CAN, TRT, US). 

Frutices grandes vel arbores parvae, plerumque trunco uno, ad 10 m alti, spinosi; ra- 
muli unius anni nirentes atro-cascanei, ramuli veteriores brunneo-cinerei; spinae plerumque 
< 2.0 cm longae, crassae, plus minusve recurvatae, in juventute brunnae nitenter. Folia 
decidua;petioli plerumque 1—2 cm longi; laminae 4— 6 cm longae, ellipticae vel lare-ellipticae, 
interdum aliquantum anguste ovatae vel rhombeae, paribus 4 venarum sccondariarum longiorum 
vel aliquando non-lobatae vel vix-lobatae vel vadosirer 2-3(-4) lobatae per latus, marginibus 
serratis, ± coriaceae, in mattiritate tenuiter appresso-pubescentes supra, subter ± glabrae 
venis exceptis, colore autumnah prunino ad carmineum vel pallide carmineum. Inflorcscenciae 
12-20 florarae, bracteolis parvis caducis glandulo-marginatis, glabrae vel raro pubescentes. 
Flores 15—20 mm diam.; hypanthium glabrum extrinsecus; lobi calycis 3 mm longi, ob- 
scure gland uloso-marginati; petala ± circulaha; stamina 10—12, antheris pallide roseis; carpelli 
et styla 3—4. Fructus ampulliformo-orbiculares, ca 10 mm diam., triste-purpurei vel nigri 
subinde in maturitate, in mense Augusto rubro-vinosi vel spadiceo-castanei, reliquiis calycis 
parvis; pyrenae 3—4, vadositer sulcatae dorsaliter, lateribus cum foveis vel interdum planis. 

Large bashes or small trees, usually single-trunked, up to 10 m tall, thorny; 
one year old twigs deep shining mahogany, older branches gray/brown; thorns 
usually < 2.0 cm long, stout, slightly recurved, shiny brown when young. 
Leaves deciduous; petioles usually 1—2 cm long; blades elliptic to broad- 



ew Crataegus 


of western North America 

Sum 18(1) 

UPGMA of taxonomic aisiance 
19 taxa 

38 characters 


chr piped — — ' 


1 ' ' 


1 20 

1 60 


Fig. 1 . Dendrogram of western North American Crcitctegi/s taxa sliowing relation.shi|TS of 
species describee! in this paper. Newly described taxa in capitals. See text lor more infor- 

elliptic, sometimes somewhat narrow ovate or rhombic, 4—6 cm long, with 
four pairs of longer secondary veins, barely lobed to shallowly 2-3(— 4) lobed, 
the margins also serrate, somewhat coriaceous, at maturity finely appressed- 
pubescent above, below ± glabrous except on the veins; autumnal color plum 
to crimson or light crimson. Inflorescences 1 2—20 flowered, with small caducous 
gland-margined bracteoles, glabrous to rarely shortly pubescent. Flowers 
15—20 mm diam.; the hypanthium externally glabrous; calyx lobes 3 mm 
long, obscurely gland-margined; petals ± circular; stamens 1 0—12, anthers 
pale pink; carpels and styles 3—4. Fruit ± amptflliform-orbicular, ca. 10 mm 
diam., dull-purple or occasionally black at full maturity, red-burgundy to 
chestnut or reddish-brown in August, calyx remnants small; nutlets 3-4, 
shallowly grooved dorsally, sides variably eroded or sometimes plane. 

Distribution. — Cratciegiis okennouii is fairly common in southern interior 
Britisli Columbia and the eastern half of northern Washington, ranging to 
Idaho and nw IMontana (Fig. 3), usually along streams or other sources of 
water. It is a tall distinctive plant, usually with a straight trunk, easily rec- 
ognized in the field both in flower and in frtiit. This new species is most 
closely related to C. Joiiglasii , sens, str., from which it differs in fruit shape 

Phipps and O'Kpnnon, New species of Crataegus from Western North America 173 

Fic;. 2. Line drawing ot C. okeinio/aiJ.B. Phipps. Inflorescence, flowers and parts from Phipps 
6959 (UWO); mfructescence, fruit and parts from Phipps 6991, 6995 aiuKYKennon (UWO). 
Scale bars = 1 cm. 


SiDA 18(1) 

Fig. 3. Distriburion map of C, okef///oi/!/ ) .\i. Fhij^ps. 

and color, flower size, color of marure leaves (often yellowish-green), habit, 
thorn length, leal shape and autumnal color of foliage. 

Crataegus okemiomi is one of a small grotip of 'hhick-frnited' (fruit ± black 
at full mattirity) species comprising series D()nglasianae(Kt\\d. ex Sarg.) Rehd. 
which I restrict to C. douglasii Lindl., its immediate relative C sz/ksclorfii 
and the new species described here. A greatly widened concept of C douit^lasn 
which includes C. saligna Greene and C. riviilarh is advocated by Welsh 
(1982) and Holmgren (1997) but in my opinion this cannot stand up due 
to many significant differences in leaf-shape, venation type, bark and thorn 
type. Moreover intermediates do not exist between saHi^na-rmdaris'^ind douglasii- 
okennonii-suksdorjii . Diffictdties for the broad concept are also emphasized 
by the fact that C. saligna (called C. doiiglasi! Linth. van diichesnensis Welsh 
by Holmgren) has 20 stamens and cream anthers, contrary to the Intermountain 
Flora. I am (JBP, 199X) therefore placing C saligna and C. rividar'is in a 
different grouping. 

Crataegus okennonii is a clearly defined segregate of C. douglasii differing 
by a considerable number of correlated characters of which thorn length 
(the shortest in series Douglasianae) and flower size (kirgest in the series) are 
the most unambiguous. The following couplet separates C. okennonii from 
the restricted interpretation of typical C. douglasii 'as, found in western Canada 
and the adjacent United States. 

Phipps and O'Kennon, New species of Crataegus from Western North America 175 

1. Usually tree-like with a straight trunk, 5-10 m tall; young twigs deep glossy 
purple-brown; thorns usually 1.5—2 cm long; flowers ca. 15—20 mm cliam.; 
fruit crimson lake or chestnut-colored in late August, becoming deep plum- 
black at maturity, not bloomy, broadly ampulliform; autumnal foliage usu- 
ally crimson, plum or pale shades of same; styles and nutlets usually 3- 
C. okennonii 

1 . Usually btishy, though often large, not conspicuously single or straight-trtmked, 
3-8 m tall; thorns often longer, to 2.5(-3) cm long; young twigs mid to 
deep glossy brown, only occasionally as dark as C okennonii; flowers often 
12-15 mm in diam.; fruit variably colored in late August though never chestnut, 
shape ellipsoid or suborbicular; autumnal foliage color often bronzy, not 
consistently ± crimson or paler; styles and nutlets 3—5 C. douglasii 

The differences between C. okennonii and a somewhat restricted C. douglasii 
are not by any means large but recognition at the specific level is also sup- 
ported by the numerical analysis (Fig. 1), the lack of intermediates among 
good quality specimens both in the field and herbarium and the consequent 
immediacy of recognition. A more elaborate comparison with C. douglasii 
in which detailed leaf shape comparisons will be used may be expected consequent 
on the publication of R. Dotterer and T.A. Dickinson's detailed biosystem- 
atic studies of C. okennonii and C. douglasii. 

It is possible that C. okennonii is the same as C. douglasii f. hadia Sarg. 
(1907), a taxon in which the fruit is described as "chestnut", certainly not 
a common color in ripening fruit of section Douglasii. However, some of 
the unripe fruit of C. okennonii seen by us certainly was the color of reddish- 
brown, highly polished, shoe leather. Piper, the original collector of forma 
hadia, variously described the fruit as brown, dark shining brown, chest- 
nut, etc. Forma badia is a taxon systematically disregarded in the floristic 
literature. Both color changes during the ripening of western North American 
Crataegus fruit and the final color of the autumnal foliage have been poorly 
understood (Phipps 1997) as taxonomic characters so we wish to record them 
here. Crataegus douglasii fruit is shown in Figures 4d and 4e. The latter, 
photographed in August, illustrates the black, pendant, ellipsoid fruit while 
the other is another British Columbia specimen showing plenty of bloom. 
Figures 4a-4c illustrate C. okennonii, plate 4a showing the late August "chestnut" 
color while Figure 4b is a later version of the same color. Figure 4c shows 
the final, near ripe color, in late September, close to that of C douglasii. The 
foliage is frequently a distinctive bright pale crimson color in late Septem- 
ber as is shown in Figure 4f. Variation in leaf form of C okennonii tends to 
fall into two classes, both illustrated in Figure 2, there being a broader, 
somewhat rhombo-deltoid, distinctly lobed form (Fig. 4a) and a more el- 
liptic or narrow-ovate, shallowly lobed shape (Fig. 4b). Crataegus okennonii 
also bears conspicuous, expanding, caducous, coral-red bud scales. 

I (J.B.P) am pleased to name this distinctive and handsome new species 


Si DA 18(1) 

Fjg. 4. Cratiicgin okmiion'ii compared with C. dniigliisii: a) C okenntDiii showing remarkable 
chestnut-coloured fruit, Kclowna, BC, 18 Aug 1995; b) C, okennonii, Form with pale foli- 
age, note ampulliform fruit, northern Okanagan, BC, 20 Aug 1995; c) C. okennonii with 
some triut close to the final purple-black color, northern Okanagan, BC, 19 Aug 1995; d) 
C. doiigLnii showing full ripe color and a high bloom, northern Okanagan, BC, 19 Aug 
1995; e) full ripe color ot C. donglasii with ellipsoidal fruit, northern Okanagan, BC, 19 
Aug 1995; f) C. okennonii showing typical cerise color of full ripe foliage; JBP in foreground; 
Palmer Lake, Washington State, 28 Sep 1993. 

after Robert (Bob) J. O'Kennon whose companionship and critical abiHties 
as a field botanist helped to make this discovery possible. It is mtriginng 
that so widely distributed a taxon, of which we are able to cite 36 speci- 
mens, has remained undetected for so long. 

Additional specimens examined; CANADA. BRITISH COLUMBIA: Northern Okanagan, 
Spallumcheen Municipality, Powerhouse Rd., M side, near Stardel Drive, alt. ca. 1350 ft, 
50°27'N, 1 19°09'W, bush, 1.5 m tall, ± erect; fruit deep plum/chestnut, ellipsoid-turbi- 
nate, 19 Aug 1995, ././3, Phtpln -"164 (UBC, UWO); Okanagan Valley, Oyama, ca. 0.5 mi 
E ofE end of spit, alt. ca. 1320 ft, 50n)7'N, 119°2rW, hedgeline on clay soil, bush 5 m 

Phiprs ani:i O'Khnnon, New species of Crataegus from Western North America 177 

tall, frtiit plum-purple, turbmate, 17 Aug 1 995 J, C. Ph!pps7158 (UWO); Castlegar, Selkirk 
College grounds, towards bottom of slope beyond beehives, scrubby area, 49° 1 9'N, 1 1 7°38'W, 
alt 950 ft, tree 6 m tall; stamens 5, anthers pale pink, buds forced til 10 May, then 10 
stamens seen, 06 May 1994 J. B. Phipjn 6928 (BRIT, CAN, DAO, TRT, UBC, UWO, US); 
Okanagan Valley, Westside Rd., ca. 9 mi W of jet. with 97 and ± due opposite Vernon, 
wooded edges on west of road, 50°1 5'N, 1 19°27'W, ca. 1200 ft, tree, 5 m tall, fruit small- 
ish, bright red-burgundy, 19 Aug 1994, Phipps.J.B. & O'Kennon, R.J. 6990 (CAN, TRT, 
UBC, UWO); Northern Okanagan, Hwy 97, E side, between Westside Rd. and Silver Creek 
Rd., alt. 1800 ft, equals JBP 6969, beautiful 7 m tree, 50°2.3'N, 119"17'W, convex light 
yellow-green foliage, fruit light red, turning to burgundy, roundish, fatter at base, glossy, 
19 Aug 1994, PhtppsJ.B. & O'Kennon. R.J. 6993 (BRIT, UWO); Salmon Arm, large haw- 
thorn pasture just N of railroad and W of central business section, alt. ca. 1900 ft, 50°43'N, 
119°16'W; bush, 4.5 m tall, fruir burgundy, locally frequent, (equals JBP 6959); 20 Aug 
1994, Phipps.J.B. & O'Kennon. RJ. 7027 (CAN, TRT, UBC, UWO); Okanagan Valley, 
Westside Rd., ca. 9 mi W of jet. with 97 and ± due opposite Vernon on E side of road, 
fenceline 50° 1 5'N, 1 19°27'W; 7 m multitrunked tree, fruit burgundy, alt. ca. 1200 ft, 19 
Aug 1994, PhippsJ.B. & O'Kennon. RJ. 6991 (BRIT, CAN, TRT, UWO); Okanagan Val- 
ley, NNE of Vernon at jet, 91197 A, scrubby hillside E side of road, 26 Sep 1993,7.6. Phipps 
andRJ. O'Kennon 6821 (BRIT, UWO). Pass Creek Rd., N of Castlegar, 1.3 mi along, mesic 
grassy slopes, alt. ca. 1800 ft, 49°19'N, 1 17°40'W, bush, 4 m tall, fruit reddish-burgundy 
(somewhat 'chestnut'-ROK), 23 Aug 1994, PhippsJ.B. & O'Kennon. RJ. 7036 (TRT, UWO); 
Castlegar, Selkirk College, path behind beehives towards river edge, open woodland-scrub, 
alt. 950 ft, 49°I9'N, 117°38'W, tree, 20 ft tall, reddish-purple foliage, dark purple, ± 
round fruit, 20 stamens, 26 Sep 1993, PhippsJ.B. & O'Kennon. RJ. 6824 (UWO); Slocan 
Valley, S of Slocan, above swamp along small road off Hwy. 6 & just N of Perry Siding 
across river, 49°40'N, 1 17°30'W, hedges and brushy slopes, alt. 1775 ft, bush 6 m tall, 
fruit red-burgundy, 22 Aug 1994, PhippsJ.B. & O'Kennon, RJ. 7047 (UWO); NNW of 
Vernon on W side Rd about 2.5-3 km SW of intersection of 97 & 97A, alt. 450 ft, 50°20'N, 
119°20'W, back of field on west side of road, bush 5 m tall, 24 Sep 1993, PhippsJ.B. & 
O'Kennon. RJ. 6785 (UWO); Okanagan Valley, Hwy. #97, ca. 2 mi S of Silver Creek Rd. 
exit, (200 m S of 'road narrows' sign), (100 m S of JBP 6968), 50°23'N, 119°17'W, alt 
1800 ft, in cut-over forest, young tree 7 m tall, 11 May 1994 J. B. Phipps 6969 (BRIT, 
TRT, UWO); Okanagan Valley, ca. 8 km NNE of center of Vernon, Pleasant Valley Rd., 
behind Vowle's residence, hedge at S boundary of Burke's PYO, 50°18'N, 119°l4'W, alt. 
380 m, dense, broad, mature hedge, dominant species, 3-7 m bush, 10 pink anthers, thorns 
long for species, 2 May 1994 J. B. Phipps 6879 (UWO); Salmon Arm, large hawthorn pas- 
ture, just N of railroad and W of central business center; 50°43'N, 119°16'W, alt. 1900 
ft, bush, 5 m tall, 10 stamens, pink anthers, 10 May 1994, J. B. Phipps 6939 (TRT, UBC, 
UWO, V); Okanagan Valley, Newport Beach campsite on W side Rd., approx. due NW of 
Vernon, end of parking lot, S end of camp, 50°20'N, 119°22'W, below trees, alt, 1140 ft, 
tree 5 m tall, erect habit, burgundy fruit, short thorns, 19 Aug 1994, Phipps. J. B. & O'Kennon, 
RJ. 6988 (BRIT, DAO, UBC, UWO, V); NNE of Vernon, S of jet. of 97 and 97 A just N 
of Baker Hogg Rd., bottom of hill, 50°21'N, 119°15'W, alt. 350 m, 4 m bush, purplish 
fruit, 23 Sep 1993, PhippsJ.B. & O'Kennon, RJ. 6821 (UWO); Okanagan Valley, E side of 
97, half mi N of intersection with 97; scrubby slopes with long cleared area, 50°21'N, 
1 19°15'W, most abundant species here, bush 6 m tall, red fruit; 19 Aug 1994, Phipps. J. B. 
& O'Kennon. R.J. 6999 (UWO); Castlegar, grassy floodplain ENE of Selkirk College, dense 
hawthorn thickets on banks of old oxbow, alt. 950 m, 49°19'N,1 17°38'W, tree, 9 m tall, 
foliage elliptical, slightly lobed, purple, fruit globose, dark purple, ? = 20 stamens, 27 Sep 
1993, Phipps. J. B. & O'Kennon. RJ. 6827 (CAN, ID, MONTU, TRT, UBC, UWO, US); 

178 SiDA 18(1) 

Norrhcrn Okana^'an, ca. 1 mi S ol Enderby, gravel driveway to NE starting at Indian Om- 
etery, alt. 1 300 ft, 5()"3 I'N, 11 9°08'W, bush, 4 m tall, fruit burgundy, only one here, 20 
Aug 1994, Phipps.J.B. &iyKeinmi. R.J. 7006 (UWO). 

U.S.A. IDAHO. Idaho Co.: US 12, ca. 10 road mi E of Kooskia, bank at back of field, 
N side of road, with large hawthorns, 46°07'N, 1 ^"50'W, alt. 1 550 ft, tree, 9 m tall, 
fruit burgundy, 26 Aug 1994, Phipps.J.B. & (rKenrioii. R.J. 7089 (UWO). MONTANA. 
Flathead Co.: Eew mi E of Columbia Falls, at dead end of Mt. Creek Rd. (off Berne l^d.), 
on fenceline opposite Shoal 330 sign, 48°22'N, 1 14°08'W, alt. 3000 ft, bush, 2 m tall, 08 
May 1994,y.Z3. Phipp.^ 693 > (UWO). Lake Co.: Route 211, ca. 4 mi S of Ronan, nr. MP 
44, at farm entrance, driveway throtigh pasture, near scenic turnoiu , alt. 3030 ft, 47°28'N, 
114°06'W, grove of hawthorns on bank S side of driveway, 20 ft apple-like tree, reddish 
foliage, black orbicular fruit, 28 Sep 1993, PhippsJ.B. & O'Kenmm. RJ. 6841 (UWO); 
Rte. 93, ca. 4 mi S of Ronan, near Milepost 44, near scenic turnout, one of group on S side 
of farm tirive, 47°28'N, 114"06'W, alt. 3030 ft, tree 5 m tall, quite large buds forced 'til 
1 1 May, stamens 10, anthers pink, (equals JBP 684 1 ), 08 May 1 994J.B. Phipps 6940 (BRIT, 
iMONTU, TRT, UWO, US). Sanders Co.: Route 200, 30 mi W of 93 on N side of road 
above Flathead R., in group of hawthorns, alt. 2840 ft, 47"I9'N, 1 14°43'W tree, 1 1 m 
tall, ± orbicular, black fruit, 29 Sep \99:\ Pb/pp.^. J. B. OiYKeinion. RJ. 6« 58 (BRIT, CAN, 
DAO, MO, UWO, V); Route 200, 50.6 km W of jet. with US 93, group of hawthorns 
above Flathead R., 47"19'N, 1 14°43'W, alt. 2840 ft, tree 9 m tall, stamens 10, anthers 
pale pink,(=JBP & RO'K 6955), 09 May 1 994,./. B. Phtpps 6955 (UWO); Route 200, 25 
mi W of 93 on N side of road above Flathead R., in long hawthorn hedge, alt. 2845 ft, 
47°19'N, 1 14°38'W, btish 6 m tall with spheroidal burgundy fruit, 29 Sep 1993, Phipps, 
J.B. & O'KeHrwn. R.J. 6857 (UWO); Route 200, exactly 42.6 km W of jet. with US 93, 
long hawthorn hedge, above Flathead River, 47°19'N, 1 14'38'W, alt. 2840 ft, (equals JBP 
& RO'K 6857), 09 May 1994, J.B, P/j/>/v 6933 (BRIT, MO, MONTU, UWO). WASH- 
INGTON. Ferry Co.: 1 Iwy. 2 1 , ca. 3 km SW of Danville, alt. ca. 500 m, 18 = 59'N, 1 1 8°3 1 'W, 
marshy roadside with trees and scrub, tree, 7 m tall, fruit burgLindy, (C. cloniilcisii very common 
here), 23 Aug 1995, ./.B. Phipp.s 7174 (UWO). Okanogan Co.: Chopaka Rd., 3 km di- 
rectly NW of Palmer Lake, open thicket, low rocky slopes, edge of floodplain, alt. 340 m, 
48°56'N, 1 19°4] 'W, tree, 5.5 m tall, fruit plum-red to burgtmdy, 28 Aug 1994, Ph/pps, 
J.B. &0'Kennon. R.J. 7105 (UBC, UWO, WS); Okanogan Valley, W side road about 7 km 
N of Tonasket, huge hawthorn stand along riverside, ca. 975 fr, 48°44'N, 1 19°25'W; bush, 
6 m tall, crimson foliage, no fruit, 3 Sep 1 993, Ph/pps. J.B. & O'Kef/i/ou. RJ. 6865 (UWO); 
Sinlahekin Valley, ca. 5 mi S of Loomis, in depression near roadside, alt. 450 m, 48°45'N, 
119°37'W, bush, 4 m tall, black fruit, 28 Aug 1994, Phipps.J.B. & O'Kenmm. RJ. 7] 09 
(UWO); ca. 6 km NE of Omak, dense hedges in valley-bottom among farm lands, alt. 250 
m, 48°27'N, 119"28'W, dense bush, 4 m tall, fruit red, changing Irom orange, 28 Aug 
1994, Phipps.J.B. & (YKmrinn. RJ. 7100 (UWO). Whitman Co.: US 195 along Spring 
Flat Creek few mi S of Colfax at MP 35.3, alt. ca. 775 m, 46"51'N, 1 17°2rW, tree, 6 m 
tall, fruit plum-red (some apiiroaching chestnut), group of same, 28 Aug 1994, Phipps. 
J.B. &(yKe)imii. RJ. 7099 (UWO); 66 Staley Rd., 1 km NE of Chambers, alt. 775 m, dry 
roadside, young tree in front yard of Harold Bough, 46''39'N, 117"! 1 'W, bush, 4 m tall, 
fruit shrivelled, 27 Aug 1994, Phipps.J.B. & (rKennon. RJ. 7096 (UWO). 

2. Crataegus okanaganensis J.B. Phipps & O'Kennon, sp. nov. (Fig. 5). 
Typi;: CANADA, British Columbia: Rough grass with hawthorns on Hwy, 97 near 
entrance to Kelowna airport, alt. 1800 ft, equals JBP 6907, bush, 4 m tall, frtiit, 
slightly dull and deep red, ± ellipsoid, 17 Aug 1994, Phipps.J.B. & (YKcnmni. RJ. 
6974 (iiolotype: UWO; isotypes: CAN, TRT, UBC, US). 

Phipps and O'Kennon, New species of Crataegus from Western North America 179 

Fig. 5. Lme drawing of C. okanaga?iensnJ.B. Phipps and O'Kennon. Inflorescence and flower 
parts from Phipps 6929 (UWO); infruccescence and fruit parts from Phipps & O'Kennon 7002 
and 6975 (UWO); and sterile shoot from Phipps 6894a (UWO). Scale bars = 1 cm. 

180 SiDA 18(1) 

Frutex, 3—6 m altiis, ramLili LiniLis anni mt-diocriter brunnci vel arrobrLinnei, in JLivencuce 
sparsim pLibescentes; spinae valiclae, 2—4 cm lon^ac, in jLivcnCLitc nitcntcr atrobrunneae, 
rccurvatae. Folia dccickia, petiolata; pctioli 1—2 cm loni^'i, sparsim pubescences, eglandulares 
vel cum glanckilis nigribus in jiiventLite (Oeston); laminae 35—6 cm longae, ovatae vel 
ovaro-rhombeae, alicjuanclo late ellipticae vel angnsre ovatae, lobis 3—4 ± acutis, marginibus 
SLibtiliter dentatis, venis 4-5 per latiis, alicjuantmn impressis stipra, in juventtite dense 
appresso-pilosis, pilis attntis m matiintate, siibter pLibescentes praecipne m venis, coriaceae, 
super nitentes praesertim in juventute, crescentes rubrae, aeneae in autumno. Inflorescentia 
panicula convexa, 10—20 florata; ramuli varie villosi, glandulo-maculati, bracteolis linearibus 
caducis glandulo-marginatis. Flotes 12—13 mm diam., conspicue ctipLililormes in |LiventLice; 
hypantliiLim externe villosum solo basale vel glabrum omnino; lobi calycis angnste trian- 
gulares, 3 — i mm longi, marginibus glandularibus, abaxiale cum pilis paticis; petala ± circularia, 
alba; stamina 10, antheris albis vel raro pallide roseis; carpelli et styli 2— 3(— 4). Fructiis 8 
mm diam., plcrumcpic urceolattis, glabcr, coccincLis in juventute sed in matuntatem coloris 
vini vel atropurpuretis (raro tere niger), si in conditione bona ctim lobis calycis longis in- 
signe erectopatentibus (seel aliquanto brevioribtis vel reflexis); pyrenae 2—3, dorsaliter stilcatae, 
latenbus ± planis, paulo pinguibus vel patilo erosis. 

Bush, 3—6 m tall; 1 year old rwigs mid to dark brown; young shoots thinly 
pubescent; thorns stout, 2—4 cm long, shiny deep brown when young, re- 
curved. Foliage deciduous, petiolate; petioles 1—2 cm long, thinly pubes- 
cent, eglandular or with small black glands when young (as at Creston, B.C.); 
blades 3.5—6 cm long, ovate to ovate-rombic, or less commonly broad el- 
liptical to narrow-ovate, with 3^ sharp lobes per side, margins finely double 
toothed, 4—5 veins per side, somewhat impressed above, densely appressed 
hairy when young above, this abrading with age, pubescent principally on 
the veins beneath, somewhat coriaceous, glossy above especially when young, 
expanding leaves reddish, the fall colour bronze. Inflorescence a domed panicle, 
10—20 flowered, the branches variably villous, gland-dotted, with cadu- 
cous linear, gland-margined bracteoles. Flowers 12—15 mm diam, notably 
cup-shaped when young; hypanthium externally villous only at biise or completely 
glabrous; calyx lobes narrow triangular, 3—4 mm long, the margins glan- 
dular, abaxially with a few hairs; petals ± circular, white; stamens 10, an- 
thers ivory or occasionally very pale pink; carpels and styles 2— 3{— 4). Fruit 
8 mm diam., generally flask-shaped, glabrous, red when young but later 
burgundy to deep purple (occasionally almost black), when in good condi- 
tion with strikingly long erecto-patent calyx lobes but these sometimes shorter 
or reflexed; nutlets 2—3, dorsally sulcate, their sides flat, a little plump, or 
shallowly eroded. 

Distribution. — Widely distributed and common from the Okanagan val- 
ley of southern British Columbia and northern Washington to northern Idaho 
(Fig. 6). This is an extraordinarily handsome hawthorn in late August and 
early September. 

Crataegus okanaganensis in similar to C. willianisii of northwest IMontana as 
illustrated by the numerical analysis (Fig. 1) but with more glossy foliage 

Phipps and O'Kennon, New species of Crataegus from Western North America 181 

Fig. 6. Distribution of C okariaf^anensis }.^. Piiipps and O'Kennon. 

of a much more coriaceous texture and the color usually yellowish-green in 
mid-season, the blades a little broader and the lobes usually less sharp, thorns 
generally stouter, different anther color, and mature fruits deep purple (Fig. 
9d) instead of sometimes blood or deep red; the young fruit is a brilliant 
red (Fig. 9e) at the stage when C. williamsii may be pale to deeper dull red. 
The color change from brilliant red (late August) to a deep dull purple (late 
September) is nearly unique among Crataegus species. In northern Idaho at 
2300 ft, C, okanaganemh flowered in 1 994 a good week before C. williamsii at 
the same altitude in northwest Montana. Otherwise identical material with 
five stamens is occasionally found. Crataegus okanaganensis was repeatedly 
collected in the 198()s in Idaho by Fred Johnson though without recognition. 
The two taxa may of ser. Purpureofructi be separated as follows: 

1. Fresh anthers white or cream; inflorescence branches thinly to moderately 
hairy; hypanthium usually glabrous; leaves coriaceous (Idaho and west). 
C. okanaganensi s 

1. Fresh anthers bright pink; inflorescence branches densely hairy; lower part 

of hypanthium densely hairy; leaves thin (nw Montana) C. williamsii 

Additional specimens examined: CANADA. BRITISH COLUMBIA: Okanagan Val- 
ley, Westside Rd. NW of Vernon, ca. 2 km S of intersection with 97, 1450 ft, hedges on 
both sides of road, 03 May 1994, Phipps. J. B. 6900 (UWO); ca. 1.2 mi E of Enderby on 
road to Mabel Lake between 200 m & 25 m W of "Autobody collision & repairs" sign, alt. 
1275 ft, hedgerow, bush, 3 m tall, fruit burgundy, 20 Aug 1994, Phipps, J. B. & O'Kennon, 
R.J. 7016 (UWO); Rough grass with hawthorns on Fiwy. 97 near entrance to Kelowna 

182 SiDA 18(1) 

airport, S side of Airport Dr., alt. ISOO ft, btish 6 m tall, fruit red, slightly deep and dull, 
± ellipsoid, 17 Aug 1994, P/npps. J. B. &0-Kc,nw,i. R.J. 697 5 (UBC, UWO); ca. 8 km NNE 
ol Vernon center at Burke's Pick-your-own on Pleasant Valley Rd., bottom of roadside, 
4()7 m; 5 m bush, dull burgundy, ellipsoid fruit, 24 Sep 199.% Phlpps. J.B. & O'KtiPwn, 
R.J. 6777 (BRIT, MONTU, UWO, WS); Shuswap Valley, just E of Enderby, across bridge, 
S side ol rd. on mound by roadside depression, alt. ca. 1 250 ft, bush, 5 m tall, 10 stamens, 
in bud, 05 May 1994, Phtpps.J.B. 6912 (UWO); Okanagan Valley, ca. 9 km N of center of 
Vernon off E side of 97, just S of jet. with 97A & just N of Baker Hogg Road, alt. .t50 m, 
hawthorn thickets to N side of long cleared ride, bush, 5 m tall, 10 white anthers, 03 May 
1994, Phipps.J.B. 6H94 (UWO); Okanagan Valley, ca. 9 km N of center of Vernon off E 
side of 97, just S of )ct. with 97A & just N of Baker Hogg Road, alt. 350 m, hawthorn 
(mainly) thickets to N side of long cleared ride, bush, 5 m tall, 10 white anthers, 03 May 
1994, Phipps.J.B. 6894a (UWO); Shuswap Valley, ca. 2.8 km E of Enderby on Kingfisher 
Rd., on hedgerow S side of road opposite house wirh wooden fence, N side, alt. I 280 ft, 
btish, 4 m tall, possibly = JBP & RO'K 6812, foliage somewhat reddish, stamens 10. cream 
anthers, 05 May 1994, Phipps.J.B. 6921 (BRIT, UWO); ca. 1.2 mi E of Enderby on road 
to Mabel Lake between 200 m & 25 m W of "Autobody collision & repairs" sign, alt. 1275 
ft, hedgerow, bush, 5 m tall, fruit reddish burgundy, 20 Aug 1994, Phipps.J.B. & (YKeinimi. 
R.J. 701 7 (UBC, UWO); Salmon Arm, large hawthorn pasture just N of railroad and W of 
central business section, alt. ca. 1900 ft, bush, 3-5 m tall, opposite building with blue 
strip, flowers very cup-shaped, anthers 10, very pale pink, 1 I May 1994, Phipps.J.B. 6966 
(BRIT, CAN, UWO); \l side of 97 A 0.5 mi N of inter.section with 97, scrubby slopes with 
long cleated area, bush, 5 m tall, scarlet fruit, 19 Aug 1994, Phipps.J.B. &0'Kenno)i. RJ. 
-^000 (UWO); Okanagan Valley, about 8 km NNE of center of Vernon, Pleasant Valley 
Rd., behind Vowle's residence, hedge at S boundary of Burke's PYO, alt. ca. 380 m, dense, 
broad mature hedge, biish, 3 m tall, nearest top of hedgerow, 10 anthers, ivory, 02 May 
1994, Phipps.J.B. 68H2 (DAO, UBC, UWO); Castlega'r, near Selkirk College,' left diand 
slope towards bottom of Rosedale Rd., alt. 970 ft, scrubby area, bush 2.5 m tall, prob. 
=JBP & RO'K 6826, stamens 10, anthers white, 06 May 1994, Phipps.J.B. 6929 (BRIT, 
TRT, UBC, UWO); Okanagan Valley, Westside Rd. NW of Vernon, ca. 2 km S of inter- 
section with 97, 1450 ft, hedges on both sides of road, 03 May 1994, Phipps.J.B. 6901 
(UWO); Okanagan Valley, west side of hwy. at entrance to Kelowna airport, alt. 1800 ft, 
open scrub, bush, 4 m tall, stamens 1 0, anthers ivory to palest pink, 05 May 1994, Phipps, 
J.B. 6907 (BRIT, TRT, UBC, UWO); West side of hwy. 21, just N of, and in view of, 
Canada Customs post, roadside/forest edge, 2200 ft, tall, 5 m bush, 27 Sep \99oJ.B. Phipps, 
J.B. & 0'Ke>i>!on. R.J. 6H32 (ID, UWO); NNW of Vernon on Wesrside Rd., ca. 2.5 km SW 
ol intersection ot 97 and 97A, hedges along roadside, back of field, W side of rd., bush, 5 
m tall with purple shrivelled fruit, 24 Sep 1993, Phipps.J.B. & O'Kennon. R.J. 6788 (C;AN, 
TRT, UBC, UWO); E side of hwy. 21, ca. 250 m S of Creston Ferry Rd. to Hood Rd., this 
species very common here, 1970 ft, dense bush, 3.5 m tall, ellipsoid, burgundy fruit, 27 
Sep I 993, Phipps.J.B. & O'Ke/i/ioii. RJ. 6831 (ID, MO, LJWO); E side of hwy. 2 I , imme- 
diately S of jet. with B(; 3, north of Oeston, alt. 1850 ft, scrubby bank, 2.5 m tall bush, 
fruit shrivelled, black, 27 Sep 1993. Phipps.J.B. & O'Kennon. R.J. 683('>(UWO); Castlegar, 
edge of Campus Rd. off Frank Beinder Rd., grassy slopes with scattered hawthorns, bush, 
3 m, Ivs sharply lobed, abundant btirgundy IrLiit, 27 Sep 1993; Phipps.J.B. & O'Kennon. 
R.J. 6826 (BRIT, UBC, UWO, US); Larry Calder property, E of 97 about 9 km NNE of 
Vernon, scrubby partly cleared hillside, 475 m, bush, 4 m, reddish foliage, fruit a good 
purple, 26 Sep 1993, Phipps.J.B. & O'Kennon. RJ. 687 9 (UWO); Kalamalka Prov. Park, S 
of Vernon, within 200 m of yellow gate, at parking lot, damjiish depression, ca. 1375 ft, 

PuiPPS A\D O'Kennon, New species of Crataegus from Western North America 183 

common, multistemmecl bush, fruit oblong-orbicular, burgundy, 24 Sep \9^y,Phipps.J.B. 

6 O'Ktminii. R.J. 6798 (BRIT, UWO); NNW of Vernon on Westside Rd., ca. 2.5-3 km 
SW of intersection -of 97 and 97A, hedges along roadside, back of field, 5 m bush with 
purple fruit, 24 Sep 1993, Phipps.J.B. & O'Kennon. R.J. 679/ (UWO); E side of hwy. 21, 
immediately S of jet. with BC3, north of Creston, scrubby banks, 1850 h, 2.5 m bush, 
leaves green turning purple, ellipsoid, burgundy fruit, 27 Sep 1993, Phipps.J.B. & O'Kennon, 
R.J. 6828 (UWO, V); N of Creston, E side of Hwy. 21, grassy banks with hawthorns on 
top, just S of junction with Hwy. 3, alt. 1850 ft, bush, 2.5 m tall, stamens 10, anthers 
white, equals JBP & RO'K 6828, 07 May 1994, PhippsJ.B. 6930 (UWO); Salmon Arm, 
large hawthorn pasture just N of railroad and west of central business section, alt. ca. 1900 
ft, bush, 3.5 m tall, fruit bright red, equals JBP 6966, 20 Aug 1994, Phipp.s.J.B. &0'Kenuon. 
R.J. 7024 (BRIT, TRT, UWO); E side of 97A 0.5 mi N of intersection with 97, scrubby 
slopes with long cleared area, young plant on south side of site, scarlet Iruit, 19 Aug 1994, 
Phipps.J.B. & O'Kennon. RJ. 7001 (UWO); E side of 97A 0.5 mi N of intersection with 
97, scrubby slopes with long cleared area, young plant on south end of site, scarlet fruit, 
19 Aug 1994, Phipps.J.B. & O'Kennon. R.J. 7002 (UWO); Hwy. 6 ca. 6 road mi W of 
Fauquier, west of Arrow Lake in Inookalin Valley, along fenceline in farmland, alfalfa fields, 
alt. 475 m, bush, 4 m tall, fruit bright red, 22 Aug 1994, Phipps.J.B. & O'Kemion. R.J. 
7045a (UBC, UWO); Okanagan Valley NNE of Vernon, Pleasant Valley Rd., hedge be- 
hind Vowle's residence, E end boundary fence with Burke's P-Y-O, alt. 380 m, bush 3 m 
tall, fruit large, plump, purple-red, hairy, with stout erect calyx lobes, 19 Aug 1994, Phipps, 

J.B. & O'Kennon. R.J. 6996 (UBC, UWO); Slocan Valley, S of Slocan, above swamp cross- 
ing river along small road off Hwy. 6 & just N of Perry Siding, hedges and brushy slopes, 
alt. 1775 ft, bush, 5 m tall, fruit bright red, 22 Aug 1994, Phipps.J.B. & O'Kennon. R.J. 
7048 (UWO); Shuswap Valley, just E of Enderby, across bridge, E side of road, 200 m N of 
gate, alt. ca. 1250 ft, level open field, growing along roadside across from cattle feeder, 4 
m tall, fresh anthers cream, 1 5 JVIay 1 995, Dowotwk. L.S. & D. 2741 (UWO); Shuswap Valley, 
just E of Enderby, across bridge, E side of road, alt. ca. 1250 ft, level open field, growing 
along roadside across from cattle feeder, 2.5 m tall, fresh anthers cream, 15 May 1995, 
Donovan. L.S. & D. 2744 (UWO); Okanagan Valley, Kelowna, service road at W side of 
airport, alt. ca. 1250 ft, fenceline, bush 3 m tall; fruit brilliant red, ellipsoid, 17 Aug 1995, 
Phipps.J.B. 7155 (UWO); Okanagan Valley, Oyama, ca. 0.5 mi E of E end of spit, alt. ca. 
1320 ft, hedgeline on clay soil, bush, 4 m tall, fruit ellipsoid, red, 17 Aug 1995, Phipps, 
J.B. 7157 (UWO); Okanagan Valley, Oyama, ca. 0.5 mi E of E end of spit, alt. ca. 1320 ft, 
hedgeline on clay soil, bush, 4 m tall, fruit ellipsoid, 17 Aug \995, Phipps. J.B. 7^9 (UWO). 
U.S.A. IDAHO. Bonner Co.: Eowcr Priest River, river bank, alt. 660 m., scarlet fruit, 

07 Aug 1897, Leiber}^.John B. 285^ (OSC); Pend Oreille R. across from Oldtown, N of 
bridge. Section 24,T56N, R6W, scrub just above high water mark, dominated by C. coliimb'iana, 
2000 ft, 07 Aug \9'66 Johnson, pyed 86126 (IDF); Eeclede, Riley Creek, 8 mi E of Priest R. 
at Pend Oreille R. (Albenai Falls Reservoir) T56N, R3W sec. 30, riparian fringe along res- 
ervoir climax forest, gentle slope, common, 12 Aug \9^'8, Johnson, P. 88079 (IDF); Pend 
Oreille R. across from Oldtown, N of bridge, T56N R6W S24, 2000 ft, shrub fringe just 
above high water mark, abundant, 07 Aug \986, Johnson. Fred 861 26 (ID (#93728); 6 mi 
E of Priest R, along Albenai Falls Reservoir in T56N, R4W sec. 34&35, riparian fringe, 
1900 ft, common, 12 Aug 1988, Johnson. P. 88082 (IDF); Section 28,T56N, R3W along 
Pend Oreille R. reservoir, roadside fringe of trees/shrubs by hayfield, many thornless trees, 
height to 15 ft, 07 Atig 1986, Johnson. P. 86124 (ID, IDF). Boundary Co.: US 95, about 
5 km S of jet. with Idaho 1 & S of turnout on right below tall conifers, alt. 2250 ft, on E 
plowed field backed by steep rocky, treed slope, dense bush, 3 m tall, stamens 10, anthers 

184 Sum 18(1) 

ivory, OH May L99 i, Plu/>/>s. J.B. 69U (ID, IJWO); US 95, ca. 5 km S of jet. with Idaho 1 
& slightly S ol turnout on ri,i^ht below tall conifers, alt. 2250 ft, on E plowed field backed 
by steep rocky, treed slope, dense bush, 3 m tall, stamens 10, 07 May 1994, Phipps. J.B. 
6933 (ID, UWO, WS); US 95 2 mi S of jet. with Idaho 1 , W side of road, roadside, 21 Sep 
199.^, Phipps. J.B. & (yKt^inin)i. R.J. 6S34 (UWO); 2.6 mi S of jct. US 95 & Hwy. 1, in 
T64N, R 1 E sec.2H (SE of C;o|->eland), roadside shrub frmge, 2200 ft, gentle slojx', 18 ft tall 
X 22 (t wide, 5 main stems 3-5, possible escape, no farms near, 12 Aug \99S, Johnson. F. 
88072 (IDF). Kootenai Co.: floodplain of Spokane R. at Coeur d'Alene, 1/4 mi W of US 
95, grassy meadow with scattered trees, no slope, open, 2000 ft, tare with C. cloz/^hisii, 10 
jun 19cS2,>/mTO;/. P.D. (IDE (#8604 IS)); Spokane River, 1/4 mi E of Washington border, 
2100 ft, among granitic rocks and sand, shrubs to 3 m, 01 Aug \9't\G, Johnson, F. 8644 (ID 
(#93729)); Coeur d'Alene, 20 Jun 1930, Ri/st.J.H. (ID), 20 Jun 19t0; floodplam of Spo- 
kane R. at Coeur d'Alene, 1/4 mi W of US 95, grassy meadow with scattered tree/shrubs, 
elev. 2000 ft, 10 Jun \9^2, Johnson. Fiecl{\r) (#93730)); lloodplain of Spokane R., at Coeur 
d'Alene, 1/-I mi W of LJS 95, grassy meadow with scattered trees & shrubs, no slope, open, 
2000 ft, 16 Jun [9^, Johnson. ED. (IDF). WASHING'EON. Okanogan Co.: Westside 
Rd. ca. 7 km N of Tonasket, 975 ft, along riverside, 7 m bush, smaller leaves, burgundy 
fruit, 30 Sep 1993, Phipps. J.B. & (VKcnnon. RJ. 6868 (UWO); Westside Rd. about 4 km 
N of Tonasket, alt. 97 5 ft, bush 5 m tall, riverside hawthorn thicket with a few poplars, 
fruit + erect, lake-red, broad-elliptic, 23 Aug 1 995, PA//;/;.i-,./.B. 7/7.5 (UWO, WS). Spo- 
kane Co.: Spokane, damp places, Siiksdorf. W.N. 919 (WS), Aug 1889. Whitman Co.: 
Pullman. P/>w. C.V. (CAS), May 1902; Westend; 1800 ft, spring 1976, May. R. 91 (RMV). 
ENGLAND: cult. Surrey, Kew, Royal Botanic Gardens; Kew #197-67, 19728; 22 May 
1985, Phipps. J.B. 5651. 

The discovery of C. ()kcinaganeiLsi.s and its clear relationship to C. wiUianisi'i 
merit the creation of a new series, as follows: 

Series Purpureofructi J.B. Phipps &0'Kennon, ser. nov. Fvpu: C. okmaganensis 
J. 11 Phipps and O'Kennon. 

Frutices, plerumc]tie grandes, vel arbores parvae; spinae mediocres in longittidine (2—4 
cmlongae). Folia plerumque ± coriacea(|uaeterC. inllicnnsii), laminae plerumqiie + rhombeae, 
(vel eliptico-oblongae vel rhombo-ovatae), plerumque lobis 3-4 brevibus patentibus pier 
latLis. Inflore-scentia 10-25 florara. Flores mediocres (10-15 mm)diam; stamina 5-10; carpelli 
et styli 2-1. Eructus coccineus, plerumcjue dare coccineus in Augusto exeunte, in maturitate 
Septembri exeunte fierens .sanguineus vel atropurpureus, orbicularis vel ampulliformis; pyrenis 
3—4, lateribus erosis vel |Tlanis. 

Bushes, ustially large, to small trees; thorns medium in length (2-4 cm). 
Leaves generally ± coriaceous (except C. willtamsii), blades usually ± rhom- 
bic (to elliptic-oblong or rhomb-ovate), usually with 3-4 short outward- 
pointing lobes per side. Inflorescences 10-25 flowered; flowers not large 
(10-15 mm diam.); stamens 5-10; carpels and sryles 2-4. Fruit red, usu- 
ally brilliant red in late August, becoming deep blood-red to dark purple 
at maturity, orbicular to flask shaped; pyrenes 3-4, sides eroded or plane. 

Distrihiituni. — A distinctive series with two known species and perhaps 
a couple of others found from the interior Pacific Northwest and southern 
British Columbia to northwestern Montana. Crataegus wiliamsii was origi- 

Phipps and O'Kennon, New species of Crataegus from Western North America 185 

nally placed in series Rotundifoltae by Eggleston but the ellipsoid fruit, long 
calyx lobes in fruit, dark-colored twigs and somewhat erose nutlets suggest 
a different affinity. Our numerical analysis (Fig. 1) indicates no relation- 
ship whatever to C. chrysocarpa in ser. Rotundifoliae. 

3. Crataegus phippsii O'Kennon, sp. nov. (Fig. 7). Type: CANADA. British 
Collmbia: Okanagan Valley, Pleasant Valley Rd., ca. 8 km NNE of Vernon center, 
bush, 4 m tall, fruit red, 24 Sep 1993, Phipps, J. B. and O'Kennon, R.J. 6780 (holo- 
type: UWO; isotyphs: BRIT, CAN, TRT, US, WS). 

Arbor vel frutex magnus, 5-6(-7) m alta, trunco uno dominante sed aliquanto cum surculis; 
ramuli unius anni atropurpureiZ-brunnei, veteres atrocinenerei; in juventute dense et breviter 
pubescens; spinae aliquantum sparsae, atrobrunneae vel nigrae, leviter recurvatae, aliquantum 
tenues, 2—4 cm. longae. Folia decidua, petiolata; petioli 1.5—2.0 cm longi, pubescentes; 
laminae 4-8 cm longae (sed ad 10 cm vel plus in blastis elongarionis), late ellipticae vel 
ovatae vel deltoideo-ovatae, lobis fere 3 ± vadositer obtusis vel late angulatis per latus, 
marginibus breve dentatis; ca. 4 (—5) venis secondariis per latus, supra breviter appresso- 
pubescentes subter breviter pubescentes praecipue in venis, aliquanrum coriaceae postea. 
Inflorescentiapaniculadepressa, 6—12 florata; ramuli dense breviter-tomentosi, bracteolati; 
bracteoli lineares glandulo-marginati. Flores 1 5—22 mm diam.; hypanthium externe breviter 
tomentosum; lobi calycis late triangulares, ca. 5 mm longi, abaxiale pubescentes praecipue 
proximale, marginibus glandulo-laciniatis; petala ± circularia; stamina 10, antheris roseis 
pallide; styli carpellique 2—4, stigmatis capitatis. Fructus 12 mm diam., breve-pubescentes, 
maturentes ab coccineo ad pruneo-purpureum vel nigrum, ± orbiculares vel late urceolati, 
lobis calycis reflexis; pyrenae 3, vadositer sulcatae dorsale, lateribus planis. 

Tree or large bush, 5— 6(— 7) m tall, with main trunk, but sometimes suckering; 
1 year old twigs dark purple-brown, older dark gray, young shoots densely 
short pubescent; thorns somewhat sparse, deep brown to black, slightly recurved, 
somewhat thin, 2-4 cm long. Foliage deciduous, petiolate; petioles 1.5- 
2.0 cm long, pubescent; blades 4—8 cm long (but on shoots of elongation 
to 10 cm or more), broadly elliptic to ovate or broad deltoid-ovate, with 
about 3 ± shallow rounded to broadly angled lobes per side, the margins 
short-dentate; about 4—5 secondary veins per side, shortly appressed hairy 
above, shortly hairy especially along the veins below, somewhat coriaceous 
later. Inflorescence a 6— I2-flowered flattened panicle, the branches densely 
short-tomentose, bracteolate, the bracteoles linear, gland-margined. Flow- 
ers 1 5-22 mm diam; hypanthium externally densely tomentose; calyx lobes 
broad-triangular, ca. 5 mm long, pubescent abaxially, especially below, the 
margins glandular— laciniate; petals ± circular; stamens 10, anthers pale pink; 
inside of hypanthial cup mostly glabrous, bristly at center; carpels and styles 
2—4, the stigmas capitate. Fruit 12 mm diam., short-hairy, ripening through 
shades of red to plum-purple to black in colour, ± orbicular to broadly flask- 
shaped, pubescent, with reflexed calyx lobes; nutlets 3, shallowly dorsally 
ridged, lateral walls ± plane. 

Distribution. — This species is not so common as the other two described 


SlDA IS(1) 

Fig. 7. Line drawing ot C. phippsii O'Kennon. Inflorescence flowers and parts from Phipps 
6891 (UWO); mfructescence, fruit and parts from Phipps 6780 and O'Kennon (UWO); two 
smaller isolated leaves from Phipps 6780 aud O'Kennon (UWO); magnified leaf (bottom left) 
from Phipps 6891 (UWO). Scale bars = 1 cm. 

Phipps and O'Kennon, New species of Crataegus from Western North America 187 

in this paper but is nevertheless widespread and has been found at fourteen 
focahties ranging from the Okanagan VaUey of southern British Columbia 
and northern Washington to the Flathead Valley of Montana (Fig. 8). We 
have never found it numerous at any site. 

Crataegus phippsii is one of the most distinctive species of hawthorn to 
have been found in North America in recent decades and does not appear 
to be closely similar to any other known in the west, a feature emphasized 
in Figure 1 . It is a large hawthorn, reminiscent of C, ?nollis, the latter a common 
species of the interior of the continent, in foliage, habit and pubescence while 
in its red (Fig. 9a) changing to purple (Fig. 9b) to black fruit it is similar to 
a number of mostly previously undescribed species of the Pacific North- 
west and British Columbia. The foliage colours attractively in autumn (Fig. 
9b, 90. The mature bark sometimes flakes off in longitudinal strips, an unusual 
feature in hawthorns (Fig. 9c). 

Crataegus phippsii was first collected by B.T. Butler (nos. 321-325) in 
1908 and these specimens were cited by Eggleston (1909) in his protologue 
for C. williamsii. The many differences from C. williamsii, however, show 
that C. phippsii in not in this affinity. That Eggleston thought it somewhat 
distinct is shown by his annotations on the sheets of "var. ovata," a name, 
however, never published. Beyond this brief glimpse of notoriety, the present 
species, very striking to the present authors, remained totally overlooked 
until our collections of recent years. 

The large, broad leaves with rounded lobes and hairiness of all parts sug- 
gests an affinity with series Molles. Superficially, it most nearly resembles 
the local species C. noelensis Sarg. of Missouri, a taxon which, however, has 
not been seen for many years. A combination of stamen number, anther color, 
foliage size and shape, and fruit color are the main characters distinguish- 
ing the species in the Molles group as shown in the following key. For con- 
venience, I am including C. greggiana in this key although it is now placed 
in set. Greggianae. 


1. Foliage small, on short shoots mainly 4 cm long. 

2. Flowers 1 — 1.5 cm diam.; stamens usually 1 C. greggiana 

2. Flowers ca. 2 cm diam.; stamens 20 C. lanuginosa 

1. Foliage larger, on short shoots generally 5-8 cm long. 

3. Anthers pink. 

4. Stamens 5-10. 

5. Fully ripe fruit orange-red; leaves ± unlobed C. noelensis 

5. Fully ripe fruit deep purple to black; leaves evidently lobed C. phippsii 

4. Stamens 20. 

6. Fruit red C. cexana 

6. Fruit yellow C. viburnifolia, C. kellogii 

3. Anthers white to cream. 


SiDA 18(1) 

Fic. 8. Distribution map of C. phippsii O'Kennon. 

7. Stamens 20 C. mollis 

7. Stamens 10 C. submoUis, sens. lat. 

Additional Specimens examined: CANADA. BRITISH COLUMBIA: Okanagan Val- 
ley, 8 km NNE of center of Vernon, Pleasant Valley Rd., behind Vowle's residence, hedge 
at S boundary of Burke's PYO, dense, broad mature hedgeline, bush 4 m, commencing 
flowering, 7-8 pale pink anthers, 02 May 1994, Phipps.J.B 6878 (BRIT, CAN, TRT, UBC, 
UWO, US); Okanagan Valley, 8 km NNE of center of Vernon, Pleasant Valley Rd., be- 
hind Vowle's residence, hedge at S boundary of Burke's PYC), dense, broad mature hedgeline, 
bush 4 m, foliage glossy, pedicels hairy, 10 pink anthers, 02 May I 994, Phipps.J.B. 6880 
(CAN, TRT, UBC, UWO); Okanagan Valley, rough, scrubby slope above dirt road NW- 
bound into O'Keefe Ranch, opposite L. & A. Cross Rd. off 97A, alt. 1290 ft a.s.I., stamens 
10, anthers light pink, equals JBP & RO'K 6803, 03 May 1994, Phipps.J.B. 6891 (UBC, 
UWO, WS); ca. 1 km N of Vernon cenrer, on dirt road N-bound into O'Keefe Ranch off 
Hwy. 97 A at L.& A. Cross Rd., dry rocky hillside, suckering clump of trees to 5 m, 25 ft 
across, bark coarse, plated, peeling, iruit purple, 25 Sep 1993, Phipps.J.B. & O'Kennon. 
R.J. 68VJJ (CAN, UWO); Pleasant Valley Rd. ca. 8 km NNE of Vernon Center, southern 
boundary hedge of Burke's PYO behind Vowles' residence, alt. ca. 380 m, dense, broad 
mature hedgeline o'^ Cratciegiis, bush 4 m tall, fruit deep red, 24 Sep 1993, Phipps, J.B. & 
O'Kennon. R.J. 6780 (BRIT, CAN, MO. UWO); ca. 5 km NNE of Enderby, on E-W rd. 
between Shuswap R. and Hwy. 9^ A, long hetlgerows, alt. ca. 1375 ft, bush, 5 m tall, truit 
red, 21 Aug 1994, Phipps.J.B. & O'Kennon. R.J. 7038 (UWO); ca. 3 km NE of Enderby on 
road to Sicamous (Hwy. 97 A), bottom of scrubby slope on east side, alt. ca. 1400 ft, clump 
of 5 m bLishes, reddish fruit, 2 1 Aug 1994, Phipps.J.B. & OKainon. RJ. 7037 (BAO, UWO); 
ca. 5 km ENE of Armstrong on E-W aligned rd. jusr ENE ot Armstrong airport, hedgerows, 
S side of road, alt. 1450 ft, bush 5 m tall, 21 Aug \994J'hipps.J.B. & O'Kennon. RJ. 704! 

Phipps and O'KiiNNON, New species of Crataegus from Western North America 189 

Fig. 9. Various pictures of C. phippsii O'Kennon and C okanagai/ensis J.h. Phipps & O'Kennon: 
a) typical reddish late August fruit color of C. phipps//, northern Okanagan, BC, 19 Aug 
1995; b) typical later September fruit color of C. phippsii with fall h)liage color beginning, 
northern Okanagan, BC, 24 Sep 1993; c) trunks of C. ph/ppsii showing shredding bark, 
shores of Palmer Lake, Washington, late Aug. 1 994; d) full ripe fruit color of C okanaganensis, 
Westside Rd., nr. Vernon, BC, 24 Sep 1 99.3; e) late August fruit color of C okanaganensis, 
nr. Kelowna, BC, 17 Aug 1995; 1) R.J. O'Kennon holding fruiting branch, Palmer Lake, 
Okanagan Co., WA, 01 Oct 1993; note fall foliage. 

190 SiDAhSd) 

(UWO); ca. 6 km N of Armstrong, branch of Deep Creek, thickecs, primarily C. doKghtsii, 
in narrow flood-plain, alt. 1450 ft, tree 4 m tall, Fruit red, 21 Aug 1994, Phipps.J.B. & 
O'Kennon. R.J. 7034^ (UWO); ca. 3 km NW of Armstrong, N-S road parallel to Schubert 
Rd. and to the W of it, E side, alone by telephone pole, alt. ca. 1400 ft a.s.l., bush 3.5 m 
tall, dark green leaves, dull reddish fruit, 2 1 Aug 1994, Phipps.J.B. & O'Kennon, R.J. 7033 

U.S.A. MONTANA. Flathead Co.: about 0.5 km along Tower Rd. off US 93, nearSW 
corner of Flathead Lake, hawthorn thicket along trail, rear of small field, alt. 2950 ft, equals 
JBP 7078, bush 7 m tall, overtopping all others, at popcorn, anthers pink, 20 May 1997, 
Phipps.J.B. & O'Kemimi. RJ. 7 591 (MONTU, UWO). Lake Co.: Hwy. 93, mipost 63.1, 
Tower Rd., (major pablillo), 2.0 mi. NW of Poison Bridge, 8-( 1 1-12)-14 flowers, 10 pink 
anthers, leaves hairy, shaggy, in green bud stage (forced), 18 May 1 995, O'Kennon, R.J. 13 1 56 
(UWO); Flathead Lake, SW border, 29 Aug 1908, Buthr. B.T. 322 (NY); Rocky Point 
Road NW of Poison at 1.7 mi N of hwy 93 and 0.3 mi NF of Jim's Road, alt. 2920 ft, 
extensive thickets near lakeshore behind new housing, bush, 5 m tall, Iruit orbicular, red- 
dening, 25 Aug 1994, PhtppsJ.B. &0'Kenmin. RJ. 7080 (BRIT, ID, MO, MONTU, UWO); 
Flathead Lake, SW border, 19 Aug 1908, Bu/hr. B.T. 323 (NY); Flathead Lane, off Tower 
Rd., olThwy 93 few mi NW of Poison, thickets along creek back of field, alt. 3050 ft, tree 

5 m tall, fruit partly deep oxblood, partly paler, 25 Aug 1994; Phipps.J.B. & O'Kennon, 
RJ. 7078 (DAO, MONTU, UWO, US); Flathead Lake, SW border, 19 Aug 1908, Butler. 
B.T. 321 (NY). WASHINGTON: Okanogan Co.: Palmer Lake, N side., open hawthorn 
thicket at upper level of pebble beach at picnic spot/cam|\sife, just E of Cdiopata Lodge, 
small tree, 6 m tall, mostly finished flowering, stamens 10, anthers pale pink in bud,=JBP 

6 RO'K 6874, 04 May 1994, Phipps.J.B. 6904 (ID, rVIO, UWO, WS); W of US 97 on 
crossroad at EUisford, in floodplain to S side of road, bush, 7 m tall, anthers 1 0, pale pink, 
equals JBP & RO'K 6876, ()4 May 1 994, Ph/pps.J.B. 6906 (US, UWO, WS); Palmer Lake, 
N side, open hawthorn thicket at upper beach level on campsite just E of Chopata Lodge, 
alt. 1 150 ft, fine tree, 7 m tall, burgundy fruit; 30 Sep 1993, Ph/pps.J.B. & O'Kennon. RJ. 
6874 (DAO, ID, MO, UWO, WS; E shores of Palmer Lake, 0.7 mi S of campsite at N end, 
thickets, alt. 355 m, bush, 4.5 m tall, deep plum-red fruit, 28 Aug 1994, Phipps.J.B. & 
O'Kennon. RJ. 7/0^' (BRIT, ID, UWO, WS); Okanogan Valley, cross-road at EUisford, alt. 
950 ft, bush 7 m tall in field in floodplain, S side of road, fruit blackish, 01 Oct 1993, 
Phipps.J.B. & O'Kennon. RJ. 6876 (UWO, WS). 


We wish to thank the National Research Council of Canada for supporting 
this research, Susan Laurie-Bourque of Hull, Quebec for the line illustra- 
tions and Antony Littlewood of the Department of Classical Studies, Tlie 
University of Western Ontario, for checking the Latin diagnoses. Peter Wood 
of Selkirk College, Castlegar is thanked for field assistance. J. B. Phipps thanks 
R.J. O'Kennon for supporting the color printing. 


Eggleston, W.W. 1909. New North American Cnitaegi. Bull. Torrey. Bot. (4ub 36:639- 

Hoi.MGRi'N, N.H. 1997. Rosaceae. In: A. Oonciuist, et al., etis. Intermountain Flora, vol. 

3A. New York Botanical Garden, Bronx. 
PiiiFPS, J.B. 199x. Taxonomic and nomenclatural problems in black and burgLmdy-truitetl 

Cviitciegns (Rosaceae) sp)ecies: C erythropodci , C. rivnLiyis and C sciligna. Sida, submitfeil. 

Phipps and O'Kknnon, New species of Crataegus from Western North America 191 

Phipps, J.B. 199H. Introduction to the red-truited hawthorns {Crataegus, Rosaceae) of western 

North America. Canad. J. Bot. 76(ui press). 
Sakcent, C.S. 1907. The black-lruited hawthorns of western Nortli America. Bot. Gaz. 

Welsh, S.L. 1987. Rosaceae. In: A Utah Flora. Great Basin Naturalist Mem. 9:520-543. 

192 SiDA 18(1) 


Annr HurcHiNcs, and Ai.ak Haxton Scott, Gillian Lfavls, and Anthony 
Cunningham. 1996. Zulu Medicinal Plants: An Inventory. (ISBN 
0-86980-893-1, pbk; ISBN 0-86980-923-7, hbk). University of Na- 
tal Press, Private Ba^^ XOI, Scottsville 3209, Republic of South Af- 
rica, in association with University of Zululand and National Botani- 
cal Institute. Exclusive distributor: hiternational Specialized Book Services, 
Inc., 5804 N.E. Hassalo Street, Portland, OR 97213-3644, U.S.A. 
(503) 287-3093; FAX (503) 280-8832. $110.00. 450 pp. 

The sLibtitk- Lindersratcs the amount ot intormation in tliis tightly organized compendium 
of information about plant u.sed by Zulu traditional healers. The authors have chosen to 
arrange their entries in phyh)genecic order to facilitate comparisons between related plants. 
As this publication deals exclusively with plants of southern Africa, the classification fol- 
lows that used in the National MerbariLim. Cieiuis numbers reflect the standards tised in 
herbaria arranged in the Englerian system with additional standards for ferns and grasses. 
These are referenced in the introduction. 

The entry lor each species includes geogra|-ihical distribution, a short ticscription, local 
names in English, Alrikaans and Zuhi followetl by documented use by Zidu healers and 
then those referring to use by other grtjtips in sotithern Africa. Physiological effects, Chemical 
constituents and biological i:)ro|ierties are added where information is available. These gaps 
will tease and stimulate further research condticred, it is hoped, under the ethical and legal 
codes iirotecting inieljecttial propert)' rights. 

There is a wealth of information not only on the medicinal uses of these plants, but by 
the social and cultural uses too. These range from love potions to charms against lightning 
and incense to invoke the goodwill of ancestors. However, the autiiors do warn readers nor 
to experiment! 

For those who do not own a copy ol Watt and Breyer-Brandwijk's 1962 edition the Me- 
dtctthil and Poisonous plants of Southern and Eastern Afriea this book is a must. For those for- 
ttinate to own a copy this will be a vakiable supplement, for at least a third of the 1032 
species mentioned here are new. 

With the worldwide renewal of interest in natural piroducts this book has a huge poten- 
tial readership. In each foreword, we arc- reminded of the value, and ephemeral nature, of 
traditional knowledge. 'Fhis book makes it less likel\' we will lose the traditional ethnobo- 
tanical knowledge of the Ztilu nMion.- 1- ion a Norris 

SiDA ISd): 192. 199S 





Department of Botany and Range Science 
Brigham Young University 
Provo. Ur 84602, U.S.A. 


USD A Forest Service Shrub Sciences Laboratory 
Provo. UT 84601. U.S.A. 


In 1874 Sereno Watson described Atriplex niittallii as a new species, citing his number 
981 collecrion, made in 1868 from northern Nevada, as a representative. His collection 
consisted ot three plants each collected from a separate population all mounted on one sheet. 
One is A. canescem^ two are the new species, A. nuttall'ii. Failure to recognize the original 
cohection and collection site of A. niittallii has resulted in assignment of the name A. niittallii 
to several different A /'r//' /ex species in western North America. However, by using cjuanti- 
tative as well as qualitative differences, A. nuttciUii is readily distinguished from its near 
relatives A. cuneata, A. falcata, A. ^ardnen, and A. tridmtata. The principal distinguishing 
features include plant size and habit, leaf dimensions, fruiting-bract size and shape, fruit- 
ing bract appendages, chromosome number, saj^onin content, flavonol content, dates of 
antliesis, and geograpiiic distribution. 


En 187-1 Sereno Watson describio Atriplex niittallii como nueva especie, citando su recoleccion 
niimero 98 1 , hecha en el norte del estado de Nevada, E.E.U.U., como representativa. Consiste 
esta recoleccion en tres plantas, de distintos lugares pero colocadas todas en un mismo pliego 
de herbario. Una de ellas es A. canescens y dos son de la nueva especie, A. nuttalli. El no 
darse cuenta los botanicos de la colecta original y su lugar de origen ha tenido como consecuencia 
la aplicacion del nombre de Atriplex niittallii a varias otras especies de Atriplex del oeste de 
Norte America. Sin embargo, usando tanto caracterfsticas cuantitativas como cualitativas, 
se distingue facilmente A. niittallii de las especies alines, A. ciineata, A. falcata, A. ganlneri, 
y A. tridentata. Las caracterfsticas de mayor utilidad son la altura de la planta, las cHmensiones 
de la hoja, la forma y tamafio de kis bracteas frucn'feras, sus apendices, el numero de cromosomas, 
contenido de saponinas, contenido de flavonoicles, fecha de antesis, y la distribucion geognifica. 


In 1 874, Sereno Warson described A /r//?/f.v nuttallii as new (Wacson 1 874) 
and listed his collection number 981, obtained in 1868 in northern Ne- 
vada, as a representative of the species. This collection consists of three plants 

SiDA 18(1): 19.0-212. 1998 

194 Sii)AlS(l) 

collected at different times in separate locations in northern Nevada, all 
mounted on one herbarium sheet (GH!) (Fig. 1), and each labeled by Watson 
as Obione carmcens Moq. The plant on the right is A. canescens (Pursh) Nutt. 
collected in June, 1868, in Unionville Valley, Pershing County, Nevada. 
The other two are A. nuttallii. The middle plant, which appears to have 
been broken and folded back, was collected in September, 1868, in Thou- 
sand Springs Valley, Elko County, Nevada. The plant on the left was col- 
lected in July 1868 in Reese Valley, Lander County, Nevada. Currently there 
are still extensive populations of these species in the valleys where Watson 
collected them. 

As shown in Figure 1, several botanists have recognized differences be- 
tween these three plants and have provided annotations for them. In 195 1 , 
Grant D. Brown annotated the plant on the right as Atnplex canescens (Pursh) 
Nutt. In 1962, C.A. Hanson annotated the plants on the left and right as 
Atriplex bonnevillensh Hanson and the central one as Atriplex falcata (Jones) 
Standley. In 1972, I.J. Bassett and C.W. Crompton annotated the plant on 
the right as Atriplex canescens, the central one as A. mittallii -vat. falcata, and 
designated on the annotation label, the left specimen as the lectotype of 
Atriplex nuttallii S. Watson. Later, McNeil et al. (1983) designated as the 
lectotype of A. nuttallii, a specimen collected in Saskatchewan, Canada, by 
Bourgeau {s.n.), in 1857, and included by Watson among 10 other collec- 
tors of A. nuttallii, in addition to Watson 981 . However, as directed in the 
International Code of Botanical Nomenclature, Article 9-9 (Greuter et al. 
1 994), — " when the material designated as type is found to belong to more 
than one taxon, a lectotype — may be designated," and, as described in 
Article 7.5 of the 1988 edition of the code (Greuter et al. 1988), "A lecto- 
type is a specimen or illustration selected from the original material to serve 
as a nomenclatural type when no holotype was designated at the time of 
publication — . When two or more specimens have been designated as types 
by the author ... the lectotype must be chosen from among them." Since 
Watson listed in the protologue, his number 981 as a representative of the 
new species, he ostensibly considered it typical and, since this collection 
consists of three specimens, one of them must be chosen as the lectotype. 
Consequently, the lectotype of Atriplex nuttallii is the specimen on the left 
(GH) as annotated by Bassett and Crompton in 1972. The designation by 
McNeil et al. of a specimen collected in Saskatchewan, Canada as the lecto- 
type of A. nuttallii, was therefore unnecessary. The illustration in Bassett et 
al. ( 1 983) of the Saskatchewan plants, is representative of a diploid A, gardneri 
(Moquin-Tandon) Hall & Clements which is common throughout south- 
ern Alberta, southern Saskatchewan and northern Montana but very differ- 
ent from A. nuttallii of northern Nevada. In his protologue, Watson de- 

Stutz and Sanderson, Atriplex nuttallii and irs near relatives 



S -'■ 

r ■ - -y Y' 


Fk,. 1. Atrip/ex ni/thiHii. The leccotypc o\' Atriplex nintnllii (Walsou 98/, upper left corner) 
collected in Reese Valley, Lander County, Nevada, July IH^iH- The central element is A. 
riiittallii collected in Thousand Springs Valley, Elko County, Nevada, September, 1868. 
The specimen on the right is Atripkxaimsmis collected in Unionville Valley, Pershing County, 
Nevada, June 1S6H. 

196 Si DA 18(1) 

scribes A. yiuttallu -^"^ being 1—2 feet high. This is the stature of the A. nuttalln 
plants in the populations he collected in northern Nevada (Table 1) but not 
A. gcirdnert plants which are mostly 6-1 2 inches tall, nor A. cciuescens plants 
which are seldom less than 30 mches in height. The Saskatchewan plant 
illustrated in Bassett et al. (1983) appears to be ca 12 inches in height. 

In the botany treatment of the King expedition (Watson 1871), Torrey 
provided identification for Watson's collections and Watson provided the 
descriptions (footnote, page 287). Referring to Watson 98 1 , Torrey identi- 
fied it as Obionecanescens Moq. and Watson applied that name to each of the 
three plants. Watson's description of the collection specifically referred to 
his number 981 and indicated that "most of the specimens have nearly or 
quite wingless fruit. Others have the bracts considerably dilated, though 
still less than is frequently the case (982)," indicating that he recognized it 
as being anomalous, quite unlike other Ohuma (Atriplex) caiiescens plants. As 
shown in Figure 2, there is considerable variation in the fruiting-bract char- 
acteristics of plants in the Reese Valley population, with none being truly 

Since Watson, following Torrey 's identification, labelled each of the three 
specimens {Watson 981), Ohione canescens, he apparently assumed the varia- 
tion he saw in the populations in northern Nevada to be representative of 
the variation displayed by Atriplex shrubs throughout western United States. 
This is confirmed in his later description of A . n/tttallii (Watson 1 874) wherein 
he lists Atriplex canescens Nuttall, not of James, Ohione canescens Moq., and 
Atriplex Gordoni Hook, as synonyms and gives its distribution as "from the 
Saskatchewan to Colorado and Northern Nevada." 

This broad interpretation by Watson appears to have been the initiation 
of a series of misconceptions concerning the geographic distribution of A. 
nuttallii that we interpret to be confined to northern Nevada and north- 
western Utah (Fig. 3). Standley (1916) listed the type locality of A. nuttallii 
as "on the denuded hills of the Missouri River, about 15 miles below the 
confluence of the White River, South Dakota." Since this is the location 
given by Nuttall (1818) as the type locality (){ Atnplex canescens (^\xi'i\\) Nutt., 
Standley clearly confused the two. Furthermore, this locality for Atriplex 
canescens {Calligonium canescens Pursh) invites correction because its origin 
given by Pursh (1814) is "in the plains of the Missouri River, near the Big 
Bend" which is several kilometers upstream from the confluence of the White 

Standley (1916) also mistakenly listed the distribution of A. nuttallii as 
"alkaline plains and hillsides, Manitoba and Saskatchewan to northern Utah, 
Colorado and western Nebnuska" without mentioning Reese Valley and Thousand 
Springs Valley in northern Nevada. 

Stutz and Sandhrson, Atriplex nutcallii and its near relatives 


Fig. 2. Fruitin^'-bracts of six randomly selected plants oi Atriplex nuttallii from a popula- 
tion in Reese Valley, ca 1 km west of Battle Mountam, Lander Co., Nevada. Each row across 
represents one individual. Bar = 15 mm. 


SlDA 18(1) 

Tic;. 3. DismbLirion ol known populations of /\/r//;/(-.v «////■/////. 

Hall and Clements (1923) likewise failed to include the type locality of 
A. niittciliii in their description of the distribution of /\ . uiittallu as "Saskatchewan 
and South Dakota to western Nebraska, New Mexico, Arizona, northwest- 
ern California, eastern Washington, and Alberta." 

Recently, Basset et al. (1983) cited the distribution of /i. uiittalliicV^ "from 
the Peace River District of Alberta to Manitoba. In the United States specimens 
have been found as far south as Colorado." Clearly this does not refer to the 
A. nuttaliii described by Watson from Nevada. 

The failure to identify the original collection site of /\. unttalli'i in all 
references has caused considerable misunderstanding of its identity and has 
resulted in the assignment of this binomial to most of the suffrutescent shrub 
species o{Atriplex in western North America. Ulbricht (1934) listed A. nuttallu 
S. Wats., A. bnxijolict Rydb., A. ciinecitci A. Nelson, A. eremicola Osterh., A. 
falcata Standley, A. i^ordomi Hook., A. nmniexkaua Standley, A. ohlanceolata 
Kydh., A. pabular is A. Nelson and A . tridentata Kuntze as synonyms oWbione 
gardneri Moq. {Atripkx gardnen [Moq.] Standley). Hall and Clements (1 923) 
treat A . buxifolia, A . cuneata, A . eremicola, A. falcata, A . gardneri, A . pabiilaris 
and A. tridentata as subspecies of nuttaliii. 

Standley (1916) listed eight species as close relatives of A. nuttaliii (Se- 
ries Nuttallianae): A . buxifolia, A . cuneata, A .falcata, A . gardneri, A. neomexicana, 
A. oblanceolata, A. pringlei Standley, and A. tridentata. As suggested by Hall 

Stutz and Sandhrson, Atriplex nuttallii and its near relatives 199 

and Clements (1923), A. pringki appears to be an insignificant variant of 
A. acanthocarpa (Torr.) S. Wats and A. ohlanceolata is non-distinguishable 
from A. gardneri. A. neomextcana is a common reoccurring hybrid between 
A. cuneata and A. conferufolia (Torr. & Frem.) S. Wats. (Hanson 1962) and 
A. bnxifolia is synonymous with A. gardneri. The remaining four species, A. 
cuneata, A. falcata, A. gardneri, and A. tndentata, were treated by Hall and 
Clements (1923) as subspecies of A. nuttallii. Since they and A. nuttallii are 
clearly distinct phenotypically (Pope 1976, Tables 1, 2, Fig. 4) and geo- 
graphically (Figs. 3, 5-8) we consider them best treated as separate species 
as proposed by Standley (1916). 

Because Watson listed A. gordoni (A. gardneri) as a synonym of A . nuttallii, 
Hanson (1962) argued that the epithet nuttallii was superfluous and hence 
illegitimate and that A . gardneri was the correct name for Watson's A. nuttallii. 
This view was later accepted by Pope (1976) Stutz (1 978), Stutz et al. (1979), 
and Welsh (1984). However, as noted by McNeil et al. (1983), the listing 
of A. gordoni (A. gardneri) as a synonym of A. nuttallii was accompanied by 
a question mark and is therefore not a legitimate synonym and, according 
to the Code, Article 55.2, note 1 , (Greuter et al. 1994) the epithet nuttallii 
is therefore not nomenclaturally superfluous. 

As discussed by McNeil et al. ( 1 983) the other synonyms of A . nuttallii 
listed by Watson: {Atriplex canescens Nuttall, and Ohione canescens Moq.), are 
also not legitimate synonyms. 


To better define A. nuttallii and its near relatives, herbarium specimens 
of the suffrutescent Atriplex species of western North America were exam- 
ined in several herbaria (BRY, CAS, GH, MO, NY, PH, RM, RSA, US, UC) 
and specimens were collected and studied from the original collection sites 
of each of the species included in the study (except A. gardneri for which 
the exact location is not known). In addition, morphological measurements, 
saponin production, flavonoid content, and cytological studies were made 
of plants in several populations of each species. The morphological mea- 
surements included fruiting-bract characteristics, plant height and width, 
and leaf length and width. Chromosome counts were obtained from plants 
in several populations of each species (Figs. 5-8). In some cases the counts 
were obtained from root tips squashed in aceto-carmine stain, but most were 
from aceto-carmine squashes of pollen-mother-cells derived from anthers 
fixed in 5% acetic acid and stored under refrigeration. Saponin content was 
determined by hemolysis of red blood-cells as described by Sanderson et al. 
(1987). Flavonoid content was determined by paper chromatography as described 
by Sanderson and Stutz (1984). 

Tahu; 1 . Plant and leaf characcensncs oi Atnl^lcx ni<t!uliu and its near relatives. Meastirements were obtained from 20 plants in each population. Diploid A. nnieata 
IS not included because ot extensive between-population variation. N = number of populations. Data = mean with coefficient of variation in parentheses. Values 
in eacli column with the same letter are not significantly different (p<.()5). 



'^^^"" f^' Heit,dit (cm) Width (cm)' Volume (dm ~)'' Len<_'rh(mm) Width (mm) 

A.withillniGx) 9 32.6(0.41)'^ 47.4(0..39)'^ 101 .5(0.94)'^ 24. 1(0. r^)"^ 4.0(0.21)'* 6.0(0.22)'^ 

A.a/iieah!(AyO 5\ 12.8(0..t])'^ 5 1.2(0. .3 .3)'^ 41.3(0.88)'^'* 20.9(0.22)'^ 8.3(0.28)'^ 2.6(0.19)'^^ 

A.l],lcjta{2^) 12 12.9(0.44)" 24.6(0.,^ 1 )'* 9.4(0.^4)" 23.7(0,58)'^ 3.6(0.41)" 6.5(0.23)'^ 

A.i^arclnendy.) 6 9.8(0.52)" 48.9(0.11)'^" 25. 2(0. 76)'^" 19.9(0.09)'^ 3.8(0.21)" 5.4(0. 1 1 )'''f^ 

A.i^m-J,m;{Ax) 8 10.9(0.59)" 49.8(0.44)-^ 42.3(1.45)'^" 19.4(0. 19)'"' 5.3(0.28)" 3.8(0.18)"™ 

A.tnJentataiGx) 1^ 23.9(0,38)'^ 41.8(0.46)^" 63.8(1 .25)'-^" 28.7(0.53)'^ 4.1(0.33)" 6.79(0.28)'^ 

' Width of A. tridentcitd does not include root sprouts. 
^ Volume = height X (width)- 




Table 2. Fruiring-bracc characteristics oi Atripkx niittaU'ii and its near relatives. Measurements were made on 20 Fruits from several randomly selected plants in 
each sampled population. Diploid A. ctineata is not included because of extensive between-population variation. N = number of populations. Data = mean with 
coefficient ol variation in parentheses. Values in each column with the same letter are not significantly different (p<.05). 


A. niittciUii{Gx) 


N Length (mm) Width (mm) 

1/w Volume (mm")'' 


No. of 
Terminal Teeth 





1.6(0.14)'^" 39.0(0.15)'^ 

2.4(0.07)'^ 3.2(0.24)R 

A.cuneata{Ax) 15 6.1(0.16)'^ 

A.falcata(2-K) 12 4.6(0. 19)^^ 

A.gardneri(,2x) 1 3.1(0.14)"= 

A.gardnertiA-y:.) 10 4.2(0.21)''^ 

A. tridentataiGx) 13 4.6(0.12)^ 

5.0(0. 22)^^ 1.3(0.14)^ 165.3(0.59)'' 1.8(0.19)*^ 3.6(0.24)^^ 

2.6(0.14)1^ 1.8(0. 10)'^ 32.0(0.48)" 3. 0(0.00)''^ l.CKO.OO)^- 

2.5(0.21)^^ 1.3(0.17)"^ 19.7(0.45)^* 1.7(0.19)"= 4.64(0.46)" 

3.0(0.31)''^° 1.5(0.12)'3c 45.1(1.12)13 1.5(0.05)^' 3.8(0.12)" 

3.9(0.16)"^: 1.2(0.15)^ 72.6(0.39)" 2.6(0.15)" 8.5(0.45)'^ 

'^ Volume = length X (width)'. 

^ Prominence of central apical tooth was scored to 3 with central tooth absent = 0, small = 1, conspicuous = 2, prominent = 3. 

■^ Number of lateral appendages was scored to 5 with no appendages = 0, numerous appendages = 5. 








Appendages'- "5 

3.9(0. 17)'^ 









Si DA 18(1) 

Fic;. 4. I'riiiting bracts ui Atr/p/cx niittallii and its near relatives. A, A. HHttaUii. B, A. trulviitcita. 
C, A. ciDiadtd. D, A. garciiien. E, A. falaita. Although there is considerable variation m the 
frtiiting bracts of plants within and between populations, tliose shown are hiirly represen- 
tative of each species. Collection sites h)r those shown are: A. unttcilUi, 1 km W ol Battle 
Mountain, Lander Co.: Nevada; A. tridisiitiitci. Rush Valley, Tooele Co.: Utah; /\. mneata, 
Navajo Mine lease site, ca 1 km SW of Farmington, San Juan Co.: New Mexico; A . y^ardneri, 
ca 10 km S oFBridger, Uinta Co.: Wyoming; A. falcutct, ca 5 km N of Ontario, Malheur 
Co.: Oregon. Bar = 15 mm. 

Stlitz and Sandhrson, Atriplex nuttallii and its near relatives 203 


Probably because they are dioecious and wind-pollinated, most of the 
shrubby species oi Atriplex in North America are unusually rich genetically 
and can often be best defined by quantitative, in addition to available qualitative, 
attributes. This is particularly true tor /\ . nuttallii and its relatives. As shown 
in Figures 3, 5-8 and Tables 1-4, although A. nuttallii and each of its near 
relatives can usually be distinguished from each other by some qualitative 
differences, when combined with differences in quantitative attributes and 
geographic distributions, they are quite distinct. 

Atriplex nuttallii S. Watson, Proc. Amer. Acad. Arts 9:116.1874. Lhcto- 
TYPi;: NEVADA: Reese Valley, Jul 1868, Watim 9H1 (GUI). 

Phenotypically, A . nuttallii appears to be most closely related to A . tndmtata. 
Although highly variable in A. nuttallii (Fig. 2), most fruiting bracts of 
both taxa have multiple, terminal, marginal teeth. Both have oblong or 
oblanceolate leaves and occupy similar habitats (primarily saline bottom- 
lands). Both are hexaploids (2/?= 54) (a few tetraploid populations of A. tridentata 
have also been found). A. nuttallii differs from A. tridentata in being taller 
statured (30-60 cm vs 10-30 cm) (Table 1), woodier, and more phenotypi- 
cally variable. Much of its variation appears to be the result of hybridiza- 
tion and subsequent introgression from other species, including A. tridentata 
with which it is often sympatric. Plants of A. tridentata do not produce sa- 
ponins whereas some plants of A. nuttallii do, some do not (Table 3). A. 
tridentata usually shows aggressive root-sprouting whereas A . nuttallii is distinctly 
caespitose with numerous (60—120), slender (1—3 mm in diameter), woody 
stems emerging from a single woody crown, 10—50 cm in diameter. Geo- 
graphically, A. tridentata is common in northern Utah, southwestern Wyo- 
ming, northwestern Colorado and northeastern Nevada (Fig. 8); A. nuttallii 
appears to be restricted to the alkaline valleys in northern Nevada and north- 
western Utah (Fig. 3). 

Atriplex cuneata A. Nelson, Bot. Gaz. 34:357.1902. Type: UTAH. Emhry 
Co.: Emery, alt, 7,000 ft, 1894, M.E.Jones 3443 (holotype: US!; isotypes: MO! NY! 

Atriplex ohlaucmlatd Rydb., Bull, Torrey Bot. Club 31:403. 1904. Type: COLORADO: 
Delta, 3 Sep 1897,././/. Onvm 4071 (holotype: US!; isotype: GH!). 

Populations of A. cuneata are restricted primarily to eastern Utah, west- 
ern Colorado and northwestern New Mexico (Fig. 5). In many places they 
constitute the dominant vegetation on thousands of acres. The common name 
of A. cuneata^ "Castle- Valley clover," connotes its value as a range plant, partly 
because it is both palatable and nutritious for livestock and wildlife and 
partly because, were it not there, there would apparently be nothing at all, 


Table 3. Cytology, saponin production, tlavonoid production, and geographic distribution ot Atrip/ex niittallti and its near relatives. The number of plants exam- 
ined is in parenthesis. 


Chrom. # 






A. nultallt! 


+ ,-(11) 


A . ciineata 


- (69) 



A . ciineata 




+ (62) 

A . fakata 

18 (115) 

+ (74) 


+ ,-(66) 

A . ii^anbien 


+ ,-(32) 


+ (31) 

A . gardnen 


+ ,-(63) 


- (40) 

A . tridetitata 

54 (257) 

- (Ill) 



Geographic Distribution 

Date of 

valleys in n NV and nw UT Jun-Jul 

e UT, w CO, nw NM Apr-May 

e UT, w CO, nw NM Apr-May 

s ID, n UT, sw WY, n NV, s OR, nw CA May-Jun 

MT, WY, AB,SK May-Jun 

MT, WY, AB, SK May-Jun 

n UT, n NV, s WY. s ID, nw CO )ul-Aug 



Stutz and Sanderson, Atriplex nuttallii and its near relatives 205 

Table 4. Key to Atriplex nuttallii and its near relatives [Series Nuttallianae of Standley (1916) con- 
sisting of species in which the plants are low-statured (less than 6 dm), suffrutescent, perennial, dio- 
ecious, shrubs. Leaves are densely furfuraceous, alternate, entire, with Kranz-type anatomy. Fruiting 
bracts are 2 — 6 mm long, 2 — 5 mm broad, usually longer than broad and usually appendaged.} 

1. Leaves linear, length more than 5 times the width 
2. Root-sprouting extensive — A. tridentata 
2. Root-spfouting none or minimal 

3. Stems 50 or more from a woody crown, each 30 — 80 cm long — A. nuttallii 
3. Stems smgle ot few, each less than 30 cm long — A. falcata 
1. Leaves ovate to oblong, length less than 5 times the width 

2. Fruiting btacts globose, 3—5 mm long with numerous flattened appendages — A. cuneata 
2. Fruiting bracts elliptical to ovoid, 1—3 mm long with few appendages — A. gardneri 

of forage value. This is conspicuous in many places where populations of A. 
cuneata terminate abruptly against clay slopes that are completely devoid of 
vegetation. It is also evident in sites where populations of A. cuneata have 
experienced large-scale "die-back" and remain empty until repopulated by 
new A. cuneata plants. 

Atriplex cuneata is mostly tetraploid but throughout its range there are 
several morphologically distinct diploid populations (Fig. 5). The variation 
present in disjunct populations of tetraploid A. cuneata is probably the result 
of introgression from these diploids, or in some cases, tetraploid A. cuneata 
may have originated polyphyletically from different diploid ancestors. 

Atriplex cuneata plants are mostly caespitose and erect with no evidence 
of root-sprouting or layering. Their fruiting-bracts are usually much larger 
than the fruiting bracts of A. gardneri (Table 2) and are covered with nu- 
merous lateral appendages (Table 2, Fig. 4). They are usually free of saponins, 
with exceptions only in populations in which introgressive hybridization 
from other species is suspected. Tetraploid A. cuneata plants test positively 
for the presence of 6-methoxy flavonols and negatively for the presence of 
3-methoxy flavonals. In contrast, diploid A. cuneata plants test negatively 
for the presence of 6-methoxy flavonols and positively for the presence of 
3-methoxy flavonols (Table 3). 

Atriplex falcata (M.E. Jones) Standley, N. Amer. Fl. 21:68. 1916. Type: 
IDAHO. Washington Co.: Weiser, alt. 2,000 ft, 7 Jul 1899, M.E. Jones s.n. {uow- 
type: RSA!; isotype: UC!). 

Populations oi A. falcata are sporadically common in southern Oregon, 
northeastern California, northern Nevada, northern Utah, southwestern 
Wyoming and southern Idaho (Fig. 6). It is mostly diploid (2;z=18), but 
occasional tetraploid populations occur (Pope 1976). 

Phenotypically, A. falcata is most easily recognized by its small stature 
(ca. 13 cm tall X 25 cm broad), small, linear leaves (ca. 24 mm long x 4 
mm wide), and small fruiting bracts (ca. 4.5 mm long X 2.5 mm wide), 


SiDA 18(1) 




A. cuneata 2x 
• A. cuneata 4x 




Fk;, 5. (icoyraphic distribution ot chromosome counts of plants of diploid (2w=18) and 
tetraploid (2w = 36) Atripkx cuneata. 

usually without appendages, and terminating in a distinct, acute, central 
apex (Tables 1,2, Fig. 4). 

The leaves of A.falcata produce abundant saponins (Table 3), a trait that 
can be useful in field identification either by blowing bubbles in a water 
emulsion of leaves or by tasting a leaf as it is chewed. When saponins are 
present, bubbles form readily in a leaf-emulsion and the leaves have a dis- 
tinctly bitter taste. Both of these tests are positive iov A.falcata leaves, negative 
for leaves of A. trulcutata plants with which A. falcata plants are sometimes 
confused. A. falcata plants are also readily distingtushed from A. trkkntata 
plants by their rooting liabit: A. triclaitata plants are vigorous root-sprouters, 
whereas A. Jalcata plants are caespitose with a single prominent taproot. 
Also, A . falcata plants produce flowers and fruits early in the spring (May- 
June) whereas A. tridentata plants flower and set fruit mostly in mid to late 
summer (July— August) (Table 3). 

Atriplex gardneri (Moq.) Standley, N. Amer. Fl. 2 1 -.GG. 1916. Typi - fSrare:-'}. 
La Platte River [date?], Gonhn 250 (hoi.otypi;: GH!, fragments). 

O/j/om- liantnen Moc]. in DC, Prod, l.rl 11. 1819. 

Populations of A. gardneri (2x, 4x) dominate thousands of acres of clay 
slopes and swales throughout much of southern and eastern Wyoming, central 
and eastern Montana, western North and South Dakota, southern Alberta, 

Stutz and SAiNDDRSON, Atriplex nuttallii and its near relatives 





• • 


% Ob 

• A. falcata 2x 
1 i A. falcata 4x 


• • 

• •• 


Fic;. 6. Geographic distribution of chromosome counts of plants of diploid (2«=18) and 
tetraploid (2w = 36) Atriplex falcata. 

and southern Saskatchewan (Fig. 7). Most populations are tetraploid but 
diploid ones are common. In southern Alberta, southern Saskatchewan and 
north-central Montana, diploid plants can usually be distinguished from 
tetraploid plants by their smaller stature, smaller, thinner leaves, finer-tex- 
tured, less woody stems and smaller fruiting bracts. However, throughout 
most of the saline deserts of Wyoming and southern Montana, diploid and 
tetraploid plants are phenotypically very similar although both show con- 
siderable phenotypic variation both within and between populations. Chro- 
mosome counts of plants in populations bordering Interstate Highway 80 
in southern Wyoming, showed a preponderance of tetraploids west of the 
Continental Divide and a preponderance of diploids to the east (Fig. 9)- 
However, no conspicuous ecological differences were evident between the 
sites occupied by tetraploids and diploids, nor was it possible to consis- 
tently distinguish diploids from tetraploids, phenotypically. This was true 
for plants in natural populations as well as garden-grown specimens. They 
are, however, clearly distinguishable by flavonoid differences: tetraploids 


SiDA 18(1) 



(D O \ 
• O 

o i 

\ o \ 

\ C> i 

I QO O gD; O 

' O'"^ o 




1 \ 


'\ o 





A. gard neri 2x 
A. gardneri 4x 

^^ 5 



Fic. 7. Geographic distribution oi chromosome cotints of plants oi' diploid {2>i=- 18) and 
tetraploid (2«= •>6) Atr/p/ex giirdiieri. 

always produce 6-mechoxy flavonols, diploids do not; diploids produce 3- 
methoxy flavonols, cetraploids do not (Table 3). Although both diploids 
and tetraploids sporadically produce saponins (Table 3), diploids are most 
olten heavy producers, tetraploids usually produce little or none. 

Atriplex gardneri plants are short-sratured, mostly 4-10 cm in height, 
(Table 1), but are often more than 80 cm in diameter. The plants are cae- 
spitose, arising from a single deep taproot but often show considerable lay- 
ering. Their fruiting-bracts are usually smaller (ca. 3^ mm) and have fewer 
lateral appendages than those of most near relatives (Table 2, Fig. 4). 

Stutz and Sanderson, Atriplex nuttallii and its near relatives 


Fig. 8. Geographic distribution of chromosome counts of plants of tetraploid (2« = 36) and 
hexaploid (2«=54) Atriplex tridentata. 

Atriplex tridentata Kuntze, Revis. Gen. PI. 2:546. 1891- T^t^e: UTAH: Corinne, 
1874, D.E.O. Kiintze 3084 (holotype: NY!). 

Two chromosome races of Atriplex tridentata have been found: tetraploid 
(2« = 36) and hexaploid (2/2=54). Hexaploids are, by far, the most common, 
occurring throughout much of western United States and in most of the 
valley bottoms in Utah and Nevada that were recently occupied by Pleis- 
tocene lakes (Fig. 8). Only three major tetraploid populations are known: 
one in Juab County, Utah, one in Eureka County, Nevada, and one in Lin- 
coln County, southwestern Wyoming (Fig. 8). Although tetraploid and hexaploid 
A. tridentata are to some extent, morphologically distinct, the differences 
do not appear to be sufficient to warrant designation as separate taxa. 

Atriplex tridentata differs from other Atriplex species in several features, 
including linear to oblong, furfuraceous leaves, fruiting bracts with con- 
spicuous terminal, marginal teeth (Fig. 4), late-flowering habit (July-Au- 
gust), and vigorous root-sprouting. A. tridentata plants are mostly herba- 
ceous above ground with a few slender stems growing erect from woody 
underground crowns and roots. 

The extent of root-sprouting in A. tridentata plants is often made con- 
spicuous by its dioecious flowering habit. By noting its flowers, the extent 
of a single male or single female plant may be easily determined. In some 
cases individual plants have been found to occupy more than 200 m^. Sometimes 


SiDA I.S(l) 






















110 115 









la E] 

EiirsicsitTi rj iiiprsi M [I , _ij IL 










ja tpjEiaj^iMjp 






j~in r:i m 











Fic;. 9. Distribution ol diploid and tetraploid populations o{ Atripkx gardneri alongside 
Interstate highway HO in southern Wyoming from the Utah border (mile 0) to near Elk 
Ivlountain Wyoming. (Chromosome counts were made on plants at each mile marker, when 
avaikible. Stippled bars = diploids, black bars = tetraploids. 

a single plant forms a complete carpet in which there are no other plants; at 
other times the root-sprouting carries a plant around and between plants of 
other species. 

Possibly because of its low saponin content (lable 3), its root-sprouting 
habit and predominantly herbaceous tissues, A. trtdentata is often regarded 
favorably by ranchers as forage for livestock (personal reports). 

Stutz and Sanderson, Atriplex nuttallii and its near relatives 211 


The authors thank Dr. Richard W. Spellenberg and Dr. Ronald L. Hartman 
for numerous helpful suggestions, Broken Hill Proprietory Minerals and 
Brigham Young University for financial assistance and the curators of the 
following herbaria for loans of specimens and access to their collections: 
BRY, CAS, GH, MO, NY, PH, RM, RSA, UC and US. 


Bassftt, I.J., C.W. Crompton, J. Mcneiu., and P.M. Taschereau. 198.1 The genus Atriplex 
(Chenopodiaceae) in Canada. Monograph No. 31. Communications Branch, Agriculture 
Canada, Ottawa. 

Greuter, W., H.M. Burdet, W.G. Chaloner, V. Demonlin, R. Grolle, D.L. Hawksworth, 
D.H. NiCHOL.s()N, P.C. Silva, F.A. Stafleu, E.G., and J. McNeil. 1988. Interna- 
tional code of botanical nomenclature. Adopted by the fourteenth International Botani- 
cal (Congress, Berlin, July-August 1987. Regnum Veg. 118. 

, F.R. Barrie, H.M. Burdet, W.G. Chaloner, V. Demoi'lin, D.L. Hawksworth, 

P.M. JORGENSEN, D.H. NicoLSEN, P.C. SiLVA, P. Trehane, and J. McNeill. 1994. Inter- 
national code of botanical nomenclatute (Tokyo Code). Adopted by the fifteenth Inter- 
national Botanical Congress, Yokohama, August-September 199.3. Regnum Veg. 131. 

Hall, H.M. and RE. Clements. 1923. The phylogcnetic method in taxonomy. Carnegie 
Institution of Washington Publication No. 326. 

Hanson, C.A. 1 962. Perennial Atriplex of Utah and the northern deserts. Thesis. Brigham 
Young University, Provo, Utah. 

McNeil, J., I.J. Ba.ssett, C.W. Crompton, and P.M. . 1983. Taxonomic and nomenclatural 
notes on Atriplex L. (Chenopodiaceae). Taxon 32: 549-556. 

Nuttall, T 1818. The genera of North American plants. Vol. 1. Philadelphia. D. Heartt. 

Pope, C.L. 1976. A phylogenetic study of the suffrutescent shrubs in the genus Atriplex. 
Ph.D. Dissertation, Brigham Young University, Provo, UT. 

PuRSii, RT. 1814. Florae Americae Septentnonalis. Vol. II. London. White, Cocchrane and 

Sanderson, S.C, and H.C. Stutz. 1984. Flavonoid aglycones of diploid and polyploid Atriplex 
amfertifolia. In: A.R. Tiedemann, E.D. McArthur, H.C. Stutz, R. Stevens, and K.L.Johnson, 
K.L. (compilers). Proc. symposium on the biology oi Atriplex and related chenopods. 
Provo, UT. 

, R.L. Pendleton, E.D. Mcarthur, and K.T. Harper. 1987. Saponin effect 

on small mammal forage preference in a planting o{ Atriplex canescens. In: F.J. Provenza, 
J.T. Flinders and E.D. McArthur (compilers), Proc. Symposium on Plant-Herbivore 
Interactions. USDA Forest Service General Technical Report INT-222, 74-77. Ogden, 

Standley, P.C. 1916. Chenopodiales. North American Flora 21(1). New York. New York 
Botanical Garden. 

Stutz, H.C. 1978. Explosive evolution of perennial Atriplex in western America. In: K.T. 
Harper and J.I. Reveal, organizers. Intermountain biogeography: A symposium. Great 
Basin Naturalist Mem.. 2:l6l-l68. 

, C.L. Pope, and S.C. Sanderson. 1979. Evolutionary sttidies ok Atriplex: Adaptive 

products from the natural hybrid SN A.tridentata _Al^ A. canesceiis. Amcr.J. Bot. 66:1181- 

212 SioA 18(1) 

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Centro de Estudios Farmacoioguvs y Botdnkos 

Consejo Nacional de Investigaciones Ckntifkas y Tecnkas 

Serrano 663, Buenos Aires, 14 14. ARGENTINA 


Strophostyles Elliot: is the only genus within the Phaseolinae with a center of distribu- 
tion in the United States. It comprises three species, namely S. hdvula (L.) Elliott, 5. umbellata 
(Willd.) Britton, and 5, leiosperma (Torrey & A. Gray) Piper, and it is considered as allied to 
Dolichopsis Hassler, a monotypic genus endemic to South America. This study analyses the 
leaf flavonoid content from the three Ssrophostyles species and Dolichopus [laraguariensis Hassler 
with the aim of examining the phylogenetic relationships among taxa. We isolated 38 gly- 
cosides based on kaempferol, quercetin and isorhamnetin. All Strophostyles species were characterized 
by the presence of isorhamnetin glycosides. However, S. leiosperma showed a distinctive 
profile while S. hdvula and S. umbellata clustered together. In contrast, D, paraguariensis 
lacked isorhamnetin-based compounds. A cladistic analysis of flavonoid plus morphologi- 
cal data supported Strophostyles monophyly and showed 5". leiosperma as the sister taxon of 
the clade S. helvula-S. umbellata. 


Strophostyles Elliott es el linico genero de las Phaseolinae con un centro de distnbucion 
en los Estados Unidos. Comprende tres especies: S. helvula (L.) Elliott, S. umbellata (Willd.) 
Britton y S. leiosperma (Torrey & A. Gray) Piper, y se considera afin a Dolichopsis Hassler, un 
genero monotfpico endemico de Sudamerica. Este estudio analiza el contenido de Oavonoides 
foliates en las tres especies de Strophostyles y en Dolichopsis paraguariensis Hassler con el objeto 
de examinar las relaciones filogeneticas entre estos taxa. Se aislaron 38 glicosidos de kaempferol, 
quercetina e isoramnetina. Todas las especies de Strophostyles se caractenzaron por la presencia 
de glicosidos de isoramnetina. Sin embargo, S. leiosperma mostro un perfil distintivo mientras 
que S. helvula and S. umbellata se agruparon juntas. Por el contrario, D. paraguariensis no 
sintetizo compuestos basados en la isoramnetina. Un analisis cladistico conjunto de los datos 
de flavonoides y caracteres morfologicos apoyo la monofilia del genero y mostro a^'. leiosperma 
como el taxon hermano del clado S. helvula-S. umbellata. 


Strophostyles EUiorr is the only genus within the Phaseolinae with a cen- 
ter of distribution in the United States. Its current taxonomic treatment 
follows the original generic concept (Elliott 1822), but it was previously 

SiDA 18(1): 213-222. 1998 

214 SiOA 1<S(1) 

associated with unrelated species and reduced to a section o^Phaseolns (de 
CandoUe 1825; Bentham 1837, 1865) until it was restored as a separate 
genus including three species (Britton & Brown 1897; Piper 1926). A set 
of characters precludes merging Strophostyles with Phaseoh/s, namely erect 
style (not coiled), lack of hooked hairs, pedicels shorter than the calyx, and 
nodes of the inflorescence somewhat swollen (Marechal et al. 1 978). More- 
over, Strophostyles plants can be recognized by their nearly asymmetric flowers 
arranged in subumbellate inflorescences, bracts and bracteoles persisting 
through seed maturation, cylindrical seeds (often pubescent), and linear pods. 

Strophostyles has also been considered as allied to Dolichopsis Hassler (Marechal 
et al. 1978; Lackey 1983). Nevertheless, both genera are easily distinguishable 
by many characters and their quite distinct geographical disrributions. The 
monotypic genus Dolichopsis^ resembles Strophostyles mainly in floral mor- 
phology (purplish corolla, keel with a not curved to somewhat curved beak, 
style slightly thickened distally, and stigma terminal oblique) and general 
appearance, but it is unlike the latter in having symmetric flowers clustered 
in elongate pseudoracemes and the unique fruit traits such as oblong, very 
flat pods with oblong seeds implanted through a very long funicle and with the 
hilum perpendictilar to the placenta. The geographic range oi Dolichopsis is Paraguay 
and Argentina, in South America, whWt Strophostyles occurs throughout eastern 
USA, eastern Canada up to south of Quebec and extreme northeastern Mexico. 

Phytochemical data on Strophostyles species are scanty and include the absence 
of both leuco-anthocyanins (Baudet 1978) and canavanine (Lackey 1977), 
and a recent report of flavonoids (Williams et al. 1995). In this study, we 
expanded on the survey of foliar flavonoids by considering a larger number 
of samples of the three Strophostyles species, i.e. S. helviila, S. mnbellata and S. 
leiospermci , and we added Dolichopsis paraguariensis for comparison. 


We analyzed the constitutive flavonoids present in the leaves of herbarium 
specimens belonging to the three Strophostyles species and Dolichopsis paraguariensis . 
Samples (100-200 mg) were powdered and extracted under reflux with 80% 
methanol (x 3). Concentrated methanoiic extracts were two-dimensionally 
chromatographed on paper (BAW/ 1 5% acetic acid). Compounds were identified 
by standard methods (Mabry et al. 1970; Markham 1982). These incltided 
complete and controlled (3 min.) acid hydrolysis, enzymatic hydrolysis (B- 
glucosidase), co-chromatography with authentic markers and UV-Vis spec- 

^Doltchupm was a genus with two species, D. paraguarienm and D, monticola (Lackey 1983, 
Lewis 1991), but recently Delgado Salinas & Lewis (1997) created the new genus Oryx'n 
where they placed D. monttadci. Tiierefore, D. paniy^/uinensis has became the tinique repre- 
sentative of the genus. 

Pelotto and Martinez, Flavonoids in Scrophoscyles and Dolichopsis species 215 

troscopy. Glucosides were separated from their galactosidic analogues by 
TLC in the appropriate system according to Budzianowski (1991). 

Plant material. — Specimens were provided by the Institute de Botanica 
Darwinion Herbarium (SI), San Isidro, and the Centro de Estudios Farmacologicos 
y Botanicos Herbarium (BACP), Buenos Aires. 

Strophostyles helvula (L.) Elliott 

U.S.A. Arkansas. Jefferson Co.: Arkansas river bottoms, 220 ft, 17 Sep 1937, Demaree 
16245 (SI). Illinois. Mc Donough Co.: Argykle Lake, near Colchester, 2 Aug \95H, Jones 
22335 (SI). Iowa. Dickinson Co.: N shore ofSpirit Lake, sand (older beach), 5 Aug 1913, 
Si>/ek 14... (number illegible) (SI). Mississippi. Harrison Co.: near the coast on sand, 6 
Jan \9'5UDewaree30675 (SI); Ship Island, P.O. Biloxi, in stabilized sand, moist, long trailing, 
Demaree 31059 (SI). Virginia. Prince George Co.: rich alluvial thicket back of sand-beach 
of James River, Jordan Point, SE Virginia, 16 Sep 1938, Feriialil & Long 9353 (SI). Un- 
hwuH loadity.movd of the Western Reserve] 15 Atig 1897, G.B. Asduroft. Berea 0. s.ii. (SI). 

Strophostyles leiosperma (Torrey & A. Gray) Piper 

U.S.A. Oklahoma: 5 mi NW of Breckenridge, 25 Jul 1941, Gephardt 747 (SI). Woods 
Co.: in waste place, hard soil, near Alva, 24 Sep \9U, Stevens 2824 (SI). Texas. Smith Co.: 
Amigo, neglected sandy field, 10-17 Aug 194'5, Moore. Jr. 995 (SI). Morris Co.: Aug 1891, 
Carleton 420 (SI). 

Strophostyles umbellata (Willd.) Bntton 

U.S.A. Virginia. Greensville Co.: dry pine and oak woods, about 1 mi N of Skipper's, 
l4-\5 ]n[ 1938, Fernalct & Long 8737 iSl). 

Dolichopsis paraguariensis Hassler 

ARGENTINA. Entre Rios: Depto. La Paz, R 1 26, desvi'o a Ombties, borde camino, 3 1 
Jan 1 98 1 , fl. azul-violaceo, Troncoso de Bi/rkart & Baagalupo 3096 (SI). PARAGUAY. Depto. 
Pte. Hayes: EstanciaLomaPyta, 23° 40'S, 59° 35 'W, 2 Apr 1974, enredadera casi rastrera, 
fl violaceas, crece en pastizal, n.v. 'kekleichetas,' Arenas 544 (BACP). Depto. Boqueron: 
Mision Santa Rosa, 21° 45'S, 61 ° 35 'W, Feb 1981, enredadera, fl. violaceas, crece en pajonal, 
n.v, 'ceihlowey'. Arenas 1726 (BACP). 

Data analysis. — A cluster analysis was performed on flavonoid data of 
15 herbarium specimens. Similarity matrix was measured using Jaccard's 
coefficient and a dendrogram was constructed applying the unweigthed pair- 
group method of arithmetic averages (UPGMA). All calculations were done 
using NT-SYS program (Rohlf 1993). 

A cladistic analysis of the flavonoid data plus a set of morphological characters 
was carried out according to the maximum parsimony principle. Outgroup 
criterion was used for character polarization. Vigna adenantha was chosen as 
the external group which flavonoid data were obtained following the methods 
above mentioned (Pelotto, unpublished manuscript). For flavonoids, char- 
acter states that occurred in the outgroup were scored as and those in the 
ingroup (D. paraguariensis and the three Strophostyles species) were scored as 
1 (see Appendix, Tables A and B). Morphological data were gathered from 
the literature and included some multistate characters that were treated as 
non-additive (see Appendix, Tables A and C). Cladograms were calculated 

-^^ SiDA 18(1) 

using the implicit enunieration routine (i.e.*) of the program Hennig86 
(Farris 1988) with all characters equally weighted. 


Chromatographic properties of the identified flavonoid glycosides and 
Its distribution in Strophostyles species and D. paragiiarienm are shown in 
Tables I and 2, respectively. All detected compounds were O-glycosides of 
flavonols with sugars attached at positions 3 and 7 of the aglicone skeleton. 
This pattern of glycosilation is very common among the Phaseolinae (Zallocchi 
& Pomilio 1994; Williams et al. 1995, Pelotto unpublished manuscript). 
All three Strophostyles species produced glycosides based on the methy- 
lated flavonol isorhamnetin plus kaempferol and quercetin. Notwithstanding 
S. helvnla and S. nmhellata showed very similar chromatographic patterns, 
while the flavonoid profile of 5. lewspmna was quite distinctive. No rhamnosides 
were detected in S. kmpmna and it only shared the presence of monoglycosides 
with the other two species. Based on a three-sample analysis within a fla- 
vonoid survey of the Phaseolinae, Williams et. al (1995) have also reported 
the occurrence of isorhamnetin glycosides in Strophostyles species and no- 
ticed the same interspecific differences. In contrast with our results, Will- 
iams and co-workers isolated fewer compounds and did not detect kaempferol 
glycosides from leaves, although they did from stems and/or flowers. 

In turn, Dolichopsisparaguariensis samples were characterized by the presence 
of kaempferol and quercetin glycosides, lacking isorhamnetin. Noticeably, 
Paraguayan samples contained only kaempferol glycosides while Argentinean 
one had kaempferol plus quercetin glycosides. However, in a previous work 
(Zallocchi et al. 1995) both kaempferol and quercetin glycosides were re- 
ported from one sample of D. pcircigiiarmisis from Paraguay, but of the eight 
flavonol glycosides the authors identified only rutin and kaempferol-3-O- 
rutinoside were also present in our .samples. These differences may be due 
to the fact that Zallocchi and co-workers analyzed a whole plant extract 
and therefore their results are difficult to compare with ours. 

After the cluster analysis S. helvnla and S. umhellata are closer to D. paraguariensis 
than to S. lewsperma (Fig. 1). This is because S. helvula and S. umhellata have 
more glycosides (based on kaempferol and quercetin) in common with D. 
paraguariensis than with S. leiosperma, even though D. paraguariensis does not 
produce isorhamnetin glycosides. 

Cladistic analysis resulted in two most parsimonious trees (length, L - 
35, consistency index, CI =91, retention index, RI =70. Fig. 2). Both cla- 
dograms support Strophostyles monophyly but differ in the depicted rela- 
tionships among Strophostyles species. One tree (Fig. 2. A) shows S. helvula 
and S. leiosperma as being sibling species, but this hypothesis needs the par- 
allel gain of the characters 19, 22 and 31 on the S. umhellata and S. helvula 
branches. The other tree (Fig. 2.B) supports the clade S. umhellata-S. helvula 

Pelotto and MARTfNEZ, Fkvonoicls in Strophostyles and Dolichopsis species 217 

Table 1. Chromatographic characteristics of the identified compounds. 


1 K-3-O-glLicoside + K-3-O-gahictoside 

2 K-7-O-glucoside + K-7-O-galactoside 

3 K-3-O-rutinoside + K-3-O-robinobioside 

4 K-3-O-diglticoside + K-3-O-digalactosidc 

5 K-3,7-0-ch^ducoside 

6 K-3-0-rLitinoside-7-0-glucoside + 

7 K-3,7-0-tnglucoside 

8 K-3,7-0-triglycoside (glu + rha + gal):|: 

9 Q-3-O-glucoside + Q-3-O-galactoside 

10 Q-7-O-glucoside + Q-7-O-galactoside 

1 1 Q-3-O-rutinoside + Q-3-0-r()binobioside 

12 Q-3-O-digliicoside + Q-3-O-digalactoside 

13 Q-3,7-0-diglucoside 

14 Q-3-0-rutinoside-7-0-glucoside + 

15 Q-3,7-0-triglucoside 

16 Q-3,7-0-triglycoside (gki + rha + gal) 

17 IR-3-O-glucoside + IR-3-O-gaiactoside 

18 IR-7-O-glucoside + IR-7-O-gahictoside 

19 IR-3-O-riitinoside + IR-3-O-robinobioside 

20 IR-3-O-diglucoside + IR-3-O-digalactoside 

21 IR-3,7-0-digIucoside 

22 lR-3-0-rutinoside-7-0-glucoside + 

23 IR-3,7-0-triglucoside 

'DP: deep purple, Y: yellow, YO; yellow-orange 
^glu: glucose, rha: rhamnose, gal: galactose 

and requires three reversions (characters 1, 9 and 17). This scenario is pref- 
erable to that portrayed on Fig. 2. A since a mutation lost is a more prob- 
able event than the homoplastic acquisition of isorhamnetm glycosides. Even 
more, if we suppose reversal of characters 1, 9 and 17 on the S. mnbellata 
branch as being a consequence of samplmg error (undersampling), the cla- 
dogram becomes shorter with only 32 steps (CI= 100, RI= 100) and the 
unique solution of a similar analysis. Thus, we consider the tree depicted 
on Figure 2.B a more plausible ingroup phylogeny. 

Flavonoid evolution shows methylation of the flavonol skeleton as an advanced 
character shared by all Strophostyles species and the absence of rhamnosides 
in S. leiosperma as an (aut)apomorphic loss. 

Morphological traits are congruent with flavonoid data. Subumbellate 
inflorescence (character 24), persistent bracts and bracteoles (character 26), 
linear, terete pods (character 27) and seed pubescence (character 28) ^\vp- 
^otxStrophostyles monophyly, and are correlated with isorhamnetin monoglycoside 
production (characters 17 and 18). Strophostyles helvula strongly resembles 


Rf(x 100) 

UV + NH3 


15% aa 




Y Y 










































DP Yo 






Y Y 



















Table 2. Glycoside disrriburion in tlie analyzed samples. Compounds are numbered according to Table 1 . K; kaempterol glycosides; Q: cjuercetin glycosides; IR: 
isorhamnetin glycosides; -r: present; -: absent. 


10 11 \2 \i 14 15 16 r 18 19 20 21 22 2t 

S. helvitla 

Jones 22335 - + + -- + --- + + -- + --- + + -- + - 

Simek 14... ++ + -- + -- + + + - - + - + + + + - - + - 

Aschroft & Berea s.n. ++ + -- + -- + + + -- + - + + + + -- + - 

Demaree 3067 5 ++ + -- + -- + + + - - + - + + + + - - + 

Demaree 1624 5 -+--- + _-- + --- + ^ + - + --- + - 

Demaree 31059 ++ + -- + -- + + + -- + - + + + + -- + - 

Fernald & Long 935 3 - + + ._ + --- + + -- + - + - + + -- + - 

S. umbellata 

Fernald & Long 87 37 - + + -- + --- + + -- + - + - + + -- + - 

S. leioiperma 

Moore Jr. 995 ++-- + --- + + -- + --- + + --^-- 

Carleton 420 +__ + + - + - + -- + + - + --i--- + ^- + 

Gephardt 747 +-- + + - + - + -- + + - + - + -- + + - + 

Stevens 2824 +-- + + - + - + -- + + - + - + -- + + - + 

D. paraguariemii 

Arenas 1726 _ + + __ + __-_____--------- 

Arenas 544 __^ + _. + -......-._. ------ 

Troncoso & Bacigalupo 3096 - + + -- + - + - + + -- + - + ------- (y^ 

— — > 


Pelotto and Martinez, Flavonoids in Strophostyles and Dolichopsis species 








— HEL-22335 

_ HEL-sn 

— HEL- 16245 

— DOL-3096 


— LEI-995 
I LEl-747 


Fk;. 1. Dendrogram of the Strophostylts and Dalichopsii specimens constructed from a simi- 
larity matrix (Jaccard's coefficient) using tiie UPGMA method. Cophenetic correlation co- 
efficient, r = 0.969 

S. umbellata, except for its more fobed leaflets and larger pods and seeds. 
Strophostyles leiosperma is rather different from the other two species because 
of the smaller flowers (character 25) arranged in more pauciflorous inflo- 
rescences and its seeds glabrous and shining at maturity (character 29). 

Similarly, both S. helvnla and S. umbellata are more widespread and northerly 
distributed, with the first species reaching Canada, while S. leiosperma has a 
more limited distribution ranging from south of the United States to the 
extreme northeastern of Mexico (Britton & Brown 1897, Marechal et al. 
1978). Species divergence at chemical and morphological level also correlates 
with their ecological features; S. helvula and S. umbellata mostly grow in 
more mesic sites, while S. leiosperma is adapted to live into more xeric habitats. 

Regarding the evolution of the growth form, overlapping this character 
onto our preferred topology suggests that annual growth would have evolved 
independently in both S. leiosperma and .S". helvula, whereas perennation would 
be the plesiomorphic state shared by D. paraguariensis and S. umbellata. 

In summary, universal occurrence of isorhamnetin-based compounds in 
Strophostyles species is a good chemical character in defining generic monophyly 
while individual glycosides are useful characters to trace species evolution. With- 
in the Phaseolinae isorhamnetin glycosides have sporadically been recorded 
in four Phaseolus species (Pelotto, unpublished manuscript) and some Vigna 
and Macroptilium species (Zallocchi & Pomilio 1994; Williams et al. 1995). 
This fact suggests that flavonol methylation has appeared several times in 
the tribe, making it a valuable phylogenetic marker at the infrageneric level. 

Beyond this contribution, flavonoid data from the related genera Oxyrhynchus 
and Oryxis are wanting for a complete view of this little group of American 
species around Vigna. 



SlDA 18(1) 


/ 3-7 


^ 18,24,26,27(2},28 

^ 2,10,25(1) 




Fic. 2. The two most parsimonious rrees (L ^ 35) ^'eneratecl using the data matrix (see 
Appendix, Table A) and Vigthi iiJenuntha as outgroup. Characters are majDped on the trees 
as Follows: solid bar - non-homoplasiotis apomorphy, clear bar = homoplasious apomorphy, 
and cross = reversal. Numbers on the right of the character symbols stand for character 
numbers (and character state). 

Pelottc) and Martinez, Flavonoids in Strophostyles and Dolichopsis species 



We thank the curators of the Institute de Botanica Darwinion Herbarium 
(SI) and the Centro de Estudios Farmacologicos y Botanicos Herbarium (B ACP) 
for providing the plant material, and CONICET for financial support. We 
are also grateful to two anonymous reviewers for suggestions that improved 
the earlier manuscript. 


Table A. Data matrix for the cladistic analysis including botii flavonoid (characters 1-15, 17-23, 
codified according to Table B) and morphological (characters 24-31, codified according to Table C) 
data sets and using S/igrui ddencintha as outgroup. 




character number 
8 9 1(1 11 12 13 li 15 n 18 19 20 21 22 2i 24 25 26 r 28 29 30 31 

y'lgna admanlki (1 > 

D. para^^uarmiili 1 1 1 1) 1 1 :' 1 

S. helviila 110 1 10 1 110 10 1 I 12 10 1 

5. iimheUata 10 10 110 10 1112 10 1 

.V. lampnma 1 1 1 1 I 1 I 1 I 1 1 1 1 1 1 1 1 I I 1 2 1 2 I 1 

Table B. Flavonoids. Character numbers are as in Table 1. Character 16 was not included because no 
hypothesis about homology can be made on a partially identified compound. 

Character states 

1. 0= absent, 1= present 

2. 0= absent, 1== present 

3. 0= present, 1= absent 

4. 0= absent, 1 = present 

5. 0= absent, 1= present 

6. 0= present, 1= absent 

7. 0= absent, 1= present 

8. 0= absent, 1= present 

9. 0= absent, 1= present 

10. 0= absent, 1= present 

11. 0= present, 1= absent 

12. 0= absent, 1= present 

13. 0= absent, 1= present 

14. 0= present, 1= absent 

15. 0= absent, 1= presenr 

17. 0= absent, 1= present 

18. 0= absent, 1= present 
19- 0= absent, 1= present 

20. 0= absent, 1= present 

21. 0= absent, 1= present 

22. 0= absent, 1 = present 

23. 0= absent, 1 = present 

Table C. Morphological characters, states and polarities. 


24. inflorescence 

25. flower size 

26. bract and bracteole 
27. pods 

28. seed coat 

29. seed pubescence 

30. hilum 

31. leaflets 

0= pseudoracemose; 1= subumbellate 

0= great (> 20 mm); 1= medium (7-15 mm); 2= small (< 7 mm) 

0= persisting no longer anthesis; 1= persisting through seed maturation 

0= linear, compressed; 1= oblong, very flat, with false cellulosic septa; 2 = 

linear, cylindrical 

0= smooth; 1= pubescent 

?= inapplicable; 0= persisting in mature seeds; 1 = absent from mature 


0= parallel to the placenta; 1= perpendicular to the placenta 

0= entire; 1 = lobed to somewhat lobed at base 

222 Sum 18(1) 


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Departaniento de Botdnica 
Universidad Autonotna Agraria "Antonio Narro" 
Buenavista, Saltillo, Coahuila 23315, MEXICO 


Rutllici jhiii/lcensis Villarreal sp. nov., from the Jimulco mountains area (southwestern 
Coahuila and northeast Durango), is described and illustrated. It is similar to R. ocadentalh 
(Gray) Tharp and Barkley, and recognized by its telatively smallet flowers, shorter corolla 
basal tube and its distribution. 


Ri/elliii jimulcensis Villarreal sp. nov. del area de las montaiias de Jimulco (suroeste de 
Coahuila y noreste de Durango), es dcscrita e ilustrada. Es similar a R. occidmtalh (Gray) 
Tharp y Barkley, y se reconoce por sus flores mas ]:iequenas, tubo basal de la corola mas 
corto y su distribucion. 

Key Words: Acanthaceae, Ruellia, Coahuila, Flora of Mexico. 

Ruellia, a tropical and subtropical genus of about 200 species is repre- 
sented in the Chihuahuan Desert Region by six species (Henrickson & Johnston, 
in press). A new species is proposed as part of the study of the Flora of Coahuila. 

Ruellia jimulcensis Villarreal, sp. nov. (Fig. 1). Typr,: MEXICO. Coahuila: Mpio. 

Torreon, Sierra de Jimulco, mina San Jose, vereda hacia la cima, 103 1.3' W, 25 ()8'N. 
Matorral con Bonetidla anomab. Agave kchitgialla. Acacia herlandien, Flourensia, Hechtia, 
SpiraeayAralia, 1 800-1850 m, 10 Ago 1994,/ A. Villarreal Q_. 1181 yM. A. Carranza. 
(holotype: MEXU; isotypes: ANSM, ENCB, TEX). 

Ruellia occidentalh (Gray) Tharp & Barkley similis sed differt flores plus brevis, corolla 
tubis plus brevis, calyx lobis longius quam corolla tubis; flores cleistogamous absens et 
differt distributio. 

Perennial herb from clustered fibrous roots; stems erect to ascending 30- 
50 cm tall, densely glandular pubescent with straight hairs about 1 mm 
long; the mternodes 4-10 cm long; leaves with petioles 6-30 mm long, 
blades broadly ovate to obovate, 3-10 cm long, 2-8 cm broad, the base 
obtuse to rounded, briefly decurrent along the petiole, apex obtuse to acute, 
the margin entire to undulate-crisped, both surfaces viscid with abundant 
glandular hairs, the dried leaves usually green-yellowish; flowers in dicha- 
sia and terminal thyrsoid panicles 5-20 cm long, to 10 cm broad, strongly 

SiDA 18(1): 223-226. 1998 


SlDA 18(1) 

Fici. 1. R/icllici jh}]//Iieiisi.s, a complete phmt 

ViLLAREAL Q., A new species of Ruellia 225 

glandular-pubescent; peduncles ascending; calyx lobes 1 5—25 mm long, linear 
attenuate, united at the very base, 1.0—1.2 mm broad at the base, unequal, 
strongly glandular pubescent; corolla funnelform, bluish-purple 30—40 mm 
long, the basal tube 8—12 mm long, the broadly campanulate throat 15— 
20 mm long, the lobes 8—14 mm long, almost as broad, erose; cleistoga- 
mous flowers absent; stamens didymanous, borne at the distal portion of 
the corolla tube, free filaments 4—10 mm long; anthers 3—4 mm long; style 
15—20 mm long; fruit ellipsoid 12—20 mm long, 3—4 mm broad, glandu- 
lar pubescent; seeds 10—12, circular to oblate, 2.0—3.0 mm long, narrowly 
winged to the apex, brownish, covered with dense apressed trichomes. 

Additional specimens examined: MEXICO. Coahuria: Mpio. de Torrcon, Sierra de Jimulco, 
proximiclacles ai ejido Trinidad, 25 08' N, 103 22' W, Matorral de Agave lechugiiiUa, Boiiteloua 
raniosa, Caesalpinia ses5iliflora,Jatropha y Yucca, ladera rocosa, 1900—2000 m, 25 Ago 1983, 
J. A. Villarreal 4387, Al.A. Carrairza y A. Rodriguez (ANSM); Sierra de Jimulco, mina San 
Jose, 25" 08' N, 103" 13' W, Matorral desertico, 1800-1850m, 11 Oct 1993, M. A. Carranza 
1951 yj. Noriega (ANSM); ca. 54 air km SSE of Torreon in canyon above Estacion Otto in 
SW side ot Sierra de Jimulco near Mina San Jose, in limestone area with Acacia, Celtis, 
Viguiera, Parthenium,Jatropha, Trixis, Fo/fq/zieria, Yucca, 25 04' N, 103 13' W, 1850 m, 12 
Sep 1980,7, Henrickson & P. Bekey IH5()4 (TEX). Durango: Mpio. de Cuencame, Sierra El 
Rosario, camino a la estacion de microondas Sapioris, carr. 49, 30 km al SE de Lerdo, 25 
24'N, 10343' W, Matorral de Acacia crassijol/a, Viguiera stenoloha, Bursera Khlen dent all , Opuntia 
imbncata y Fouquieria splendem, 1750-1800 m, 16 Ago 1991, J-A. Villarreal 6243 y M.A. 
Carranza (ANSM); Estacion de microondas Sapioris, ca. 30 km SW of Gomez Palacio on 
Hwy to Durango 25" 24'30" N, 103" 43' W, matorral desertico microfilo, 1400-1 500 m, 
25 Mar 1973, M.C. Johnston, T.L. Weridt & F. Chiang 10409 (TEX); just SSE of Estacon 
Microondas Sapioris, about 20 km NW of Estacion Chocolate, 25" 25' N, 103" 43' W, 1450- 
1500 m, 14 Aug 1973, M.C.Johnston. T.L. Wendt. F Chiang &J. Henrickson 12210 (TEX); 
Microondas Sapioris, along cobblestone road which departs from Hwy 40 N of Estacion 
Chocolate, ca 15 air mi (25 km) W-SW of Torreon, 25" 25' N, 103" 42' W, Teconia stans. 
Agave lechuguilla. Euphorbia antisyphilitica 'And d'wevsc c-Act'i, 1300 m, 30Jul 1991, AI. Mayfield, 
A. Hempel & A.Jack 1093 (TEX); Mexico Hwy 40, 25 mi SW of Lerdo, 6 nov 1964, D. 
Flyr 251 (TEX); Mpio. de Lerdo, 4 mi southwest ol Chocolate, route 31, growing benerh 
Prosopis on clay flat in valley, 23 Jul 1 958, D.S. Correll & I.M.Johnston 20008 (TEX); ca. 4 
mi SW ot Ciudad Lerdo along Hwy 40 to Zacatecas, on limestone hillside with Larrea, 
Jatropha, Acacia, Opuntia, Agaveetc.,2'i" 3V N, 103 32' W, 1200 m, 2 I Sep 1978,/. Henrickson 
&E. Lee 17474 (TEX). 

Ruellia ji??iulcensis is found on sandy hillsides and dry canyons at eleva- 
tions of 1200— 2()()() m, in xeric shrublands at the complex of mountains 
near Sierra de Jimulco and Sierra del Rosario. 

The new species has leaf blades broadly ovate, obtuse to rounded at the 
base as R. occidentalis and often with strongly stipitate glandular trichomes 
covering the stems and inflorescence. It differs in its flowers 3—4 cm long, 
the basal tube of the corolla 8—12 mm long, the calyx lobes longer than the 
basal tube, the cleistogamous flowers unknown and its distribution, as marked 
in the key below. Tharp and Barkley (1949) gave the name R. occidentalis 

226 SiDA 18(1) 

WAt. ferrisae to a specimen from the mountains near Monterrey, Nuevo Leon 
which is recognized by Henrickson and Johnston (in press) by this name 
for the specimens described here. The type specimen from the Monterrey 
area has oblong-obovate leaf blades, lacks stipitate glands on stem and leaves 
and flower dimensions that better fit with R. nudiflora, a common species 
in Monterrey area. The populations of the proposed species grow allopatri- 
cally in an isolated area at the southwestern corner of the Chihuahuan Desert 
Region while most of the related species are distributed in the eastern Si- 
erra Madre Oriental (Turner 1991)- 

The three species of Ruellia previously mentioned can be separated by 
the following key: 

i . Leaf blades ovate to elliptic-obovate, mostly less than 3 cm wide, sparsely to 

moderately pubescent; lower internodes pubertilent to glabrate R. nudiflora 

1. Leaf blades broadly ovate to subdeltoid, 4-7(— 9) cm wide, usually densely 

pubescent; lower internodes with abundant stipitate glands 2 

2. Flowers (chasmogamous)4.5-5.5{— 6.5) cm long; basal tube of the corolla 
2.5—3.5 cm long, longer than the calyx lobes; cleistogamous flowers (smaller 
than the chasmogamous) frecuently located in the lower nodes; se Texas, 

ne Mexico (e Coah, ne N.L., Tamps, ne Vcr.) R. occiclentalis 

2. Flowers 3.5—4.0 cm long; basal tube of the corolla H— 1 2 mm long, shorter 
than the calyx lobes; cleistogamous flowers absent; sw Coahuila and ne 
Durango R . j i mulcensis 


I thank Tom Wendt for the loan of specimens for revision from TEX/LL. 
The illustration was prepared by Cuauhtemoc Gonzalez de Leon. 


Henrickson, J. and M.C.Johnston, (m press). A flora of the Chihuahuan Desert region. 

{Ruellia). Los Angeles, California. 2:1 151-1 155. 
Tharp, B.C. and F. Barki.i;y. 1949. Genus Rtidlui in Texas. Amer. Midland Nat. 42: 1-86. 
Turner, B.L. 1991 . Texas species oiRi/ellia (Acanthaceae). Phytologia 7 1 :281-299. 





Jardin Botdnico, U.N. A.M. 

A. P. 70-614. Del Coyoacdn 

04310 Mexico. D.F., MEXICO 


Depto. de Botdnica, Imtituto de Biologia, U.N. A.M. 

A. P. 70-233. Del. Coyoacdn 

04310 Mexico, D.F., MEXICO 


Se describe e ilustrs. Agave gracilis de los estados de Guerrero y Oaxaca, Mexico. La especie 
pertenece al grupo Striatae Baker, del subgenero Littaea (Tagliabue) Baker, y muestra si- 
militudes con A. dasylirioides ^diCohi & Bouche. 


Agave gracilis from Guerrero and Oaxaca, Mexico, is described and iUusrrated. The spe- 
cies is a member of group Striatae Baker, subgenus Littaea (Tagliabue) Baker. It is similar 
to A. dasylirioide.s JdLQohl & Bouche. 

Las exploraciones botanicas recientes realizadas en los estados de Guerrero 
y Oaxaca, Mexico, revelaron la existencia de una nueva especie de Agave, pertene- 
ciente al subgenero Littaea (Tagliabue) Baker, grupo Striatae Baker, de la 
familia Agavaceae. El grupo Striatae es endemico de Mexico y los cinco taxa 
que lo conforman se caracterizan por presentar hojas estriadas, lineares, con 
el margen serrulado, flores campanuladas o cilindricas con el tubo bien desarroUado 
y ovario sin cuello que se proyecta hacia el interior del tubo (Gentry 1 982). 

Agave gracilis Garcia-Mend. & E. Martinez, sp. nov. (Fig. 1). Tipo: MEXICO. 
GuHRRniio: Municipio de Tlapa, El Salado, 8 km al N de Tlapa, camino a Huamuxtitlan, 
selva baja caducifolia, 990 m, 16 Nov 1982 (fls), £. Martimz et al. 2659 (iiolotipo: 
MEXIJ; isoTiPo.s: BRIT, ENCB, K, MO). 

Plantae perennes, caespitosae. Folia 40-60 x 0.4—0.9 cm, linearia, striata, margine subtiliter 
dcnticLilata, glauca vel viridi-glauca. Flores campanulati 2— 2.2(— 2.5) cm longi, tubo 3—4 
mm longo, 5—7 mm lato; filamenta 2.7—3 cm, longa, in apice tubi inserta; ovaritim collo 
carens, in tubi interiorem projeccum; capsulae globosae 9—10 x 8—9 mm. 

Plantas perennes, cespitosas, con troncos rastreros de hasta 1 m de largo; 
rosetas hemisfericas, compactas, 50—80 cm de diametro, 50—60 cm de alto. 

SiDA 18(1): 227-230. 1998 


SmA 18(1) 

Fig. 1 . Agave gracilis, a) planca complt-ra con infloresccnt ia, b) bracteas superiores del pedunculo, 
c) hoja, d) detalle del margen dc la hoja, e) flores pareatlas en la inflorescencia, f) flordisectada, 
g) capSLilas, h) scmillas. Ilustrat ton basada en los especi'mcnes E. Mitrti'iiuz et uL, 2639 y A. 
Gam'd-Mi'iuioZLi y L. de hi Rosa 6"^ OH. 

Garci'a-Mendoza and Martinez, Una nueva especie de Agave de Mexico 229 

Hojas mas de 100 por roseta, 40-70 cm de largo, 0.4-0.9 cm de ancho en 
la parte media, ensanchandose en la base hasta 1-1.5 cm, lineares, estriadas, 
planas, flexibles, glaucas o verde-glaucas, subcoriaceas; margen finamente 
denticulado, amarillento; espina 3— 6(— 8) mm de largo, debil, de color pardo- 
rojizo. Inflorescencia de 1.8-2 m de largo, erecta o ligeramente inclinada, 
espiga en el cuarto superior o mitad superior del pedunculo, bracteas del 
pedunculo 5-15 cm de largo, 2-4 mm de ancho en la base, disminuyendo 
en tamaiio hacia el apice, lineares, pardas, debiles, sin espina o esta apenas 
marcada; bracteas florales 2-3.5 cm de largo, 1-2 mm de ancho, lineares, 
pardas, persistentes, mas iargas que las flores. Flores 2-2.2(-2.5) cm de largo, 
campanuladas, verdes, con el apice de los tepalos rojizo-oscuros; pedicelos 
1 mm de largo, alargandose a 2 mm durante la fructificacion; ovario 7—10 
mm de largo, 2-4 mm de ancho, cilindrico, sin cuello, glabro o glabrescente, 
penetrando ligeramente en el tubo del perianto, este de 3-4 mm de largo, 
5-7 mm de ancho; tepalos 0.9-1 ■ 1 cm de largo, 2.5-3.5(-4.5) mm de ancho, 
oblongos; fdamentos 2.7-3 cm de largo, insertos en el apice del tubo; anteras 
7-9 mm de largo, 1 mm de ancho, centricas, rojizas. Capsulas 9-10 mm de 
largo, 8-9 mm de ancho, globosas, pardo-oscuras, con el perianto, estambres 
y estilo de la flor persistentes. Semillas 3-3.5 mm de largo, 2-2.5 mm de 
ancho, engrosadas en el lado curvo, negras. 

Paratipos; MEXICO. Guerrero: Municipio de Atlixtac, 1 km al O de Santa Isabel, 30 
km al E de Chilapa, carr. a Tlapa, 25 Nov 1989 (A), J.L. Contreras 2651, 2632 (FCME); 
Municipio de Tlapa, rio El Salado en su union con el rio Tlapaneco, 8 km al N de Tlapa, 
carr. a Huamuxtitlan, 7 May 1997 (fr), A. Garci'a-Mendoza y L. de la Rosa 6508, 6311, 
6514 (ENCB, MEXU); Municipio de Zumpango del Rio, Xocohite, Xochipala, 23 Abr 
1993 (fr), A. Gomez s.n. (MEXU). Oaxaca: Distrito de Huajuapan, El Boqueron, cafion del 
n'o Mixceco, 5 km al N de Tonala, 13 Ago 1993, A. Garcia-Mendozay F. Palma 581 1 (MEXU). 

La especie aqui descrita se locaHza en la cuenca interinedia del rfo Balsas, 
en los estados de Guerrero y Oaxaca. Seguramente su distribucion se extiende 
hacia otros afluentes del rio y posiblemente mas alia, siendo quiza, la planta 
senalada por Ullrich (1990) para el cerro Yucuyu en el Distrito de Tlaxiaco, 
Oaxaca. Agave gracilis crece en laderas y barrancas abruptas, sobre suelos derivados 
de rocas calizas, entre los 850 y 1300 m snm. Habita preferentemente en 
las selvas bajas caducifolias y su ecotonia con los bosques de Quercus-Pinus. 
Algunos de los generos frecuentes con los que convive son: Acacia, Agave, 
Bur sera, Euphorbia, Hechtia, Ficus, F orchhammeria, Neobuxbaumia y Pachycereus. 

Agave gracilis presenta similitudes morfologicas con Agave dasylirioides 
Jacobi & Bouche, especie de la que se diferencia por su habito cespitoso, 
roseta mas reducida en tamaho, hojas mas cortas y angostas, inflorescencia 
erecta o ligeramente inclinada y flores mas pequehas; las capsulas globosas 
y de tamano mas reducido contrastan con las capsulas oblongas y mas grandes 

230 SiDA 18(1) 

'['ahi_a 1 . ('ompuracion lIc algunas caracai'i'sricas niorfolof^icas entrc Agari' i^ycnilis y /i- cliisyHrio'ulei. 

(^anicrer Agiive gnu'ilts Agave dcisylirinida 

Flantas cespitosas solitarias 

'IVonco hasta Im, rastrcro hasca 1.5 m, rasrrero 

Diamecro de la roseca 50-80 cm 100-200 cm 

'nimano de hojas 40-70 X 0.4-0.9 cm (40_)6()-l 00 x 2-.^. 8 cm 

C^olor de hojas ghuicas o vcrdc-^i^diiLicas verdes o verde-i,daLicas 

Iiitloresccncia 1.8-2 m, erecta o (1.5-)2-2.6 m, arqucatla 

ligeramenre inclinatia 

I.ongiCLid de florcs 2-2.2(-2.5) cm (2.6-)3-,t.5 cm 

Tubo de la tlor 3-4 X 5-^ mm (8-)l()-15 X (8-)12-]6 mm 

Tamano de Hlamcntos 2.7-3 cm 35-5 cm 

Inscrcioii de filamenros apice del tubo mitad del rubo 

Capsulas 9-10 X 8-9 mm, giobosa.s (I0-) 15-20 X 6-9 mm, obiongas 

Semillas 3-V5 X 2-2.5 mm 3 — i X 2.5-^ mm 

de A. clcLsylirioides; asi mismo, las semillas son de tamano menor en A. gra- 
cilis. Una comparacion de los caracteres mas sobresalientes de ambas especies 
se presenta en la Tabla 1 . Las medidas de A. dasylirioides se tomaron en parte 
de Gentry (1982) y se ampliaron de acuerdo con observaciones propias. Ambas 
especies habitan en la cuenca del rio Balsas, pero A. dasylirioides ocupa la 
parte alta, mas htlmeda y fria, entre los 1900 y 2500 m snm, sobre laderas 
con suelos derivados de rocas volcanicas, en bosques de Querms-Pin//s con 
Alniis. Arbutus, Clethra, Cremnophila, Garrya, Hechtia y Salvia. La epoca de 
floracion de las dos especies se presenta en los meses de octubre y noviembre. 
El epfteto especifico se refiere al caracter esbelto de la planta. 


Agradecemos la revision del manuscrito a Raquel Galvan, Susan Verhoek, 
Wendy Hodgson y Fernando Chiang. La descripcion latina fue hecha por el 
ultimo autor. La ilustracion es aportacion de Albino Luna. 


Gentry, H.S. I9H2. A^^avcs ol Continental North America. University o( Arizona Pres.s, 

Ui.LRiOi, B. 1990. Em neuer Standort Hir Agave chisylirionlcs ]i}i<:oh\ & Bouciie in Oaxaca. 

Kakreen Sukk. A 1 (H): l64- 1 66. 



Department of Botany 

University of Texas 

Austin. TX78713, U.S.A. 


In preparation for a forthcoming treatment of tlie Apocynaceae of Mexico, a routine examination 
of herbarium specimens revealed the following new species, MandevUla pririgki ].¥^. Wil- 
liams, sp. nov. The new species is distinct from other members of its alliance (sect. Torosae) 
in having subsessile leaves and larger peduncles and flowers. In addition, Mandevilla apocynifoUa 
(A. Gray) Woodson is here presented to be a synonym of Al. foliosa (Miill. Arg.) Hemsl. 


En la preparacion de un proximo tratamiento tie las Apocynaceae de Mexico, un examen 
de rutina de los especimenes de herbario revelo la nueva especie siguienre, Mandevilla pringlei 
J.K. Williams, sp. nov. La nueva especie se distingue de otros miembros de su alianza (sect. 
Torosae) por tener hojas subsesiles, y pedunculos y flores mas grandes. Ademas, Mandevilla 
apocynifoUa (A. Gray) Woodson se presenta aquf como un sinonimo de M. foliosa (Miill. 
Arg.) Hemsl. 

Key words: Apocynaceae, Mandevilla, }'A\%c<.^, Mexico 

Mandevilla pringlei J. K. Willliams Sp. nov., (Fig. 1). Type: MEXICO. Jalisco: 
Ri'o Blanco, near Guadalajara, 12 Jul 1902, C.G. Pringle 1 1337 (hoeotype: US!). 

Mandevilla foliosa (Miill. Arg.) Hemsl. affinis sed (oliis subsessilis petiolis 2-3 mm longis 
(vice 10-18 mm in M. foliosa), pedunculis longioribus (7-10 cm vice 0.3-1.2 cm) lobis 
coroUae longioribus latioribusque (12-15 X 10-14 mm vice 5-8 X 2-4 mm) plantis 
suffrutescentibus (vice lignosibus ramificantibusque). 

Suffrutescent herbs to 0.4 m tall, stems pubescent. Leaves 3-7 cm long, 
opposite, subsessile, pubescent; petioles 2-3 mm long; blades 3-7 cm long, 
1.8-2.5 cm wide at middle, ovate-elliptic, apex acute, base sub-cordate, 
with 2-4 glands at apex of petiole on upper side. Inflorescence lateral, in- 
determinate, racemose, with 2-10 flowers; peduncles 7-10 cm long, pu- 
bescent; bracts 4-6 mm long, narrowly triangular to lanceolate, straight; 
pedicels 10-17 mm long, pubescent, occasionally twisted. Sepals 5, basally 
fused, 5-7 mm long, ca 0.5 mm wide, narrowly triangular to lanceolate, 
straight, pubescent. Corolla salverform, yellow; tube 11-18 mm long, constricted 
at mouth, lower half internally glabrous, upper half internally pubescent; 
lobes 12-15 mm long, 10-14 mm wide, obovate, acuminate, occasionally 
equal in length to the tube but always greater than half its length, spread- 

SiDA 18(1): 231-235. 1998 


SlDA 18(1) 



s:Ars or ;ai.:s:o 

'-— ■■» /■— .. 

Fic;. 1. H()loty|ie of AL/Wfr///i//^;7;/,(,'/(7 J.K. Williams. 

Williams, A new species of Mandevilla 233 

ing, pubescent. Stamens 3—4 mm long; filaments ca 0.5 mm long, pubes- 
cent, straight; anthers ca 3 mm long, bases sagittate with blunt lobes. Pis- 
tils 7—11 mm long; style 5—7 mm long, glabrous; ovary ovoid, ca 1 mm 
long, glabrous; pistil head pentagonal, 2—3 mm long. Nectaries 5, as long 
as to slightly shorter than ovary. Mature follicles unknown, immature fol- 
licles fused at apex, pubescent. 

Additional specimens examined: MEXICO. Jalisco: Mpio. iVIazamirIa, Fraccionamiento 
Los Cazos, to the S of Mazamitla, 9 Jul 1995, Machuca 7351 (TEX); Cerro Viejo, S face, 
above Zapotitan de Hidalgo, a village 1 mi N of Hwy iVIEX 15, ca 25 mi due S, or 45 road 
mi from Guadalajara, base of mountain in open meadow just E of trail, alt 1890 m, 27 Jun 
1956, D. P. Gregory & G. Eiten 225 (P, SMU); Huejotitan, Jul 1912, Digi/et s.u. (P). 

Distribution. — Mandevilla pringlei is known only from five collections 
made from the Pine-Oak forest near Guadalajara, Jalisco, Mexico. 

The new species honors Cyrus Guernsey Pringle (1838-1911) promi- 
nent collector of the Mexican flora, and whose collections led to many a 
new species in the Apocynaceae, and other families. 

All specimens are with flowers and buds, Machuca 7331 is also with fruit 
but the fruits are extremely immature, only 5 mm in length. The closest 
relative o{ Mandevilla pringlei, M. foliosa (MiLill. Arg.) Hemsl., has an aver- 
age follicle length of 10 cm. 

It should be noted that according to Pringle's diary (Davis 1936, p. 199) 
on the days prior to and after his collections made on July 22, 1902 he was 
collecting numbers in the 8600 series. This, however, is in contrast to the 
type collection of M, pringlei which is numbered 1 1357. The diary, how- 
ever, does specifically state that Pringle was collecting along the Ri'o Blanco, 
the type locality of AI. pringlei, on July 22, 1902. In the numerical listing 
of Pringle's collections (ibid) the number 11337 is used twice. One listing 
is for E. apocynifolia A. Gray (= M. foliosa see below) the second is for an 
unidentified species of Asclepiadaceae. In the absence of a thorough expla- 
nation of Pringle's numerical system it is reasonable to assume that the collection 
number of the above type specimen is simply an error in numbering. 

To date the most taxonomically thorough investigation o(I\Aanckvilla remains 
Woodson's (1933) revision. Infrageneric relations are at present moderately 
unclear, however, Woodson provided both subgeneric and sectional divi- 
sions which remain undisputed. 

I^landevilla pringlei is a member of subgenus Mandevilla (as evidenced by 
the lack of glands along the midrib of the upper surface of the leaves) where 
it relates to section Torosae, evidenced by its suffrutescent habit and non- 
twinning stems (Woodson 1933). Mandevilla pringlei is most closely related 
to M. foliosa, sharing with it an erect habit (opposed to the trailing habits of 
M. karwinskii (Miill. Arg.) Hemsl. and M. torosa (Jacq.) Woodson) and hav- 


Sum 18(1) 

Fic;. 2. Comparision of flowers. A. Mandevilla foliosa (Miill. Arg.) Hemsl. (V. Fmik 2766, 
TEX). B. Mandevilla prmglei].K. Williams (P. Gregory & G. Etten 225, SMU). Bhick bar 
represents i cm. 

ing leaves with an average length greater than 5 cm (vs. 2—5 cm of AI, mexicana 
(Mlill. Arg.) Woodson). Mandevilla pringlei differs from M. foliosa in a number 
of floral and habit characters that are contrasted below: 

1. Branching shrubs to 1.5 m tall; leaves petiolate, petioles 10—18 mm long; 
peduncles 0.3—1 -2 cm long; corolla lobes 5-8 ( 1 0) mm long, 2—4 mm wide, 
up to btit not exceeding half the length ot corolla tube (Fig. 2a); stems, leaf 
blades, inflorescence, and fruit glabrous or pubescent, but midrib of leaves 
always pubescent; midrib hairs linear lanceolate O.f 5— 0.3 rnm long (Fig. 
3a); throughout Mexico M. foliosa 

1 . Sutfrutescent herbs ro 0.4 m tall; leaves subsessile, petioles 2—3 mm long; 
peduncles 7—10 cm long; corolla lobes 12—15 mm long, 10—14 mm wide, 
half or more the length of the corolla tube (Fig. 2b); stems, leaves, inflores- 
cence and fruit pubescent; midrib hairs triangular, 0.1—0.15 mm long (Fig. 
3b); Jalisco IVI. pringlei 

In the most recent keys to the species oi Mandevilla (Woodson 1933, 1938), 
M. pringlei keys out to M. apocynifolia (A. Gray) Woodson (BASfONYM: Echites 
apocynifolia). In fact, Woodson (1933) cites the type of Al. pringlei as a specimen 
of AI. apocynifolia. An examination of an isotype of Al, apocynifolia {Palmer 
734; holotype: GH; isotype: NY!), however, reveals that it is a synonym of 
M. foliosa {Ghiebreght s.n.\ holotype: G n.v., photo-holotype F! MO!). Al- 
though the type of AI. apocynifolia is without flowers, it is evident that the 
sheet represents a specimen of AI, filiosa due to its branching, petiolate leaves, 
glabrous stems and fruits, and linear-lanceolate midrib hairs. Mandevilla 
foliosa has both glabrous and pubescent-stemmed populations scattered 
throughout its range. However, in the region of Jalisco where the type of 
AI, apocynifolia was collected the populations have consistently glabrous stems 
and fruits. 

Williams, A new species of Mandevilla 



Fig. 3. Scanning electron micrographs of Mandevilla midribs of leaf undersurface showing 
the length and shape of midrib hairs. A. Mandevilla foliosa (Mull. Arg.) Hemsl. {R. King &. 
T. Soderstrom 4632, TEX). B. Mandevilla prmglei J. K. Williams (Machuca 7351, TEX). White 
bar on both photos represents 0.1 mm. Microphotographs made by theauthor using a Phillips 
515 SEM (Cell Resource Center, University of Texas Austin). 


I would like to thank Paul Fryxell for providing the Latin diagnosis and 
to him and Billie Turner for reviewing the original manuscript. Thanks is 
also extended to the curators of F, MO, NY, P, SMU, TEX and US for al- 
lowing me the opportunity to observe their specimens. I would also like to 
credit the staff of the Cell Research Center (University of Texas at Austin) 
for allowing me access to their scanning electron microscope. 


Davis, H.B. 1936. Life and work of Cyrus Gtiernsey Pringle. Univ. Vermont, Burlington. 
Woodson, R.E., Jr. 1933. Mandevilla: In; Studies in the Apocynaceae IV. Ann. Missouri 

Bot. Card. 20:645-777. 
1938. Mandevilla. In: N.L. Britton et al., eds. N. Amer. Fl. 29:645-777. 

236 SiDAlS(l) 


Fautin, Daphni; Gail, Douglas J. Futuyma, and Frances C. Jamls, eds. 
1996. Annual Review of Ecology and Systematics. Volume 27. 
(ISBN 0-8243-1427-1 , hbk). Annual Reviews Inc., 4139 El Camino 
Way, P.O. Box 10139, Palo Alto, CA 94303-0139. $52.00. 648 pp. 

There is quite a mix ot articles reviewing' tlie fields of ecolo^^y and systematics for J 996. A 
total of 20 articles are presented followed by a Subject Index, Cumulative Index of Con- 
tributing Authors, and a Cumulative Index of Chapter Titles, Volumes 23-27. The con- 
tents are as loUows: Early history and progress of women ecologists: Emphasis upon re- 
search contriburions. Forest Canopies: Methods, hypotheses, and future directions. Extinction 
by hybridization and introgression. Evolutionary significance of resource polymorphisms 
in fish, amphibians, and birds. Management of the Spotted Owl: A case history in conser- 
vation biology. Trouble on oiled waters: lessons from the Exxon Valdez Oil Spill. Evolu- 
tionary significance ot local genetic differentiation. Rates of molecular evolution: Phylo- 
genetic issues and applications. Herbivory and plant defenses in tropical forests. Mechanisms 
creating community structure across a freshwater habitat gradient. Natural freezing sur- 
vival in ainmals. Demograjihic and genetic models in conservation biology. Gene trees, 
species trees, and systematics. Incidence and consequences of inherited environmental ef- 
fects. Recruitment and the local dynamics of open marine populations. When does mor- 
phology matter.-' Adaptive evolution of photoreceptors and visual pigments in vertebrates. 
Microbial biodiversity: Domains and Kingdoms. The geographic range: Size, shape, boundaries, 
and internal structure. 

Jami:.s Stijbhendii;ck, Stephan L. Hatch, and Charles H. Butji-rfield. 1997. 
North American Range Plants. Fifth edition. (ISBN 0-8032-9243- 
0, pbk). University of Nebraska Press, 312 Nortii l4th Street, Lin- 
coln, NE 68588-0484. (402)472-3581. $25.00. 501 pp. Illustrated. 

"The 200 species in this book were selected because of their abundance, desirability, or 
noxious properties. The list of plants was developed over a nearly 40-year period by coaches 
of range plant identification teams and faculty from the colleges and universities with range 
management programs. The formal list is now the Master Plant List for the International 
Range Plant Identification Contest sponsored by the Society for Range Management (1839 
York Street, Denver, CO 80206)." Each species is treated with the following information: 
Tribe, species, common name, life span, origin, and season, followed by more detailed in- 
formation on Inflorescence Characteristics, Vegetative Characteristics, Growth Character- 
istics, Forage Value, and Habitat. An dlustration and range map are provided for each plant. 

SiDA 18(1): 236. 1998 



Department of Botany 

University of Texas 

Austin. TX 78713, U.S.A. 


In preparation tor a forthcoming treatment oi the Apocynaceae of Mexico, it was discov- 
ered that Echites holosericea of Sesse & Mo^. has priority over the species Mandevilla sertnligera 
Wocxlson and Al. syrinx Woodson, accordingly Mandevilla holosericea (Sesse & Mog.) J.K. 
Williams comb. nov. is here proposed. 


En la preparacion de un proximo traramiento de las Apocynaceae de Mexico, se descubrio 
que Echites holosericea Sesse & Mog. tiene prioridad sobre Mandei'illa sertidigera Woodson y 
M. syrinx Woodson, como consecuencia se propone Mandevilla holosericea (Sesse & Mog.) 
J.K. Williams comb. nov. 

Mandevilla holosericea (Sesse & Mog.)J.K. Williams, comb. nov. Basionym: 
Echites holosericea Sesse & Mog., Fl. Mex. (ed. 2) 45. 1894. Type: MEXICO. MiCHOACAN: 
"Ahualulci sylvis," Oct 1787-1795, ^&v.«'<& Mogino 3073 (holotypr: MA!; isotype: F!). 

Mandevilla syrinx Woodson, Ann. Missouri Bot. Gard. 19:53. 1932. Typi:: MEXICO. 

Jalesco: barranca of Tequila, 8 Oct 1893, Pringle 5422 (Hc:iE(yrYPE: MO n.v.). 
Mandevilla sertuligera Woodson, Ann. Missouri Bot. Gard. 19:383. 1932. Type: MEXICO. 

Michoac;an: rocky hills, Coru Station, 23 Jan 1907, Pringle 1 3890 (hoeotype: US!). 

Representative specimens. MEXICO. Jalisco: baranca near Guadalajara, Jun 1886, Palmer 
98 (US). MEXICO: San Lucas, district Temascaltepec, 28 Nov 1933, Hvnton 5274{VSy, 
San Lticas, district Temascaltepec, 26 Dec 1933, Hinton 7 176 (US); Dist. Temascaltepec, 
Volcan, 1410 m, 9 Aug 1932, Hinton 1295 (US), MICHOACAN: rock-fields, Coru Sta- 
tion, 15 Oct 1904, Pringle 13106 (TEX, US-2 specimens). MORELOS: lava beds near Cuernavaca, 
500 ft, 23 Jun 1896, Pr/ngle 6329 (US). OAXACA: 5 km E of Tamazulapan, road to Chilapa, 
district of Teposcolula, 1800 m, 7 Jun 1985, Mendoza et al. 1467 (NY). 

Sesse and Mo^ino (1887, 1894) described 19 species of Echites and the 
types for a majority of these names have not been located. In preparation of 
a forthcoming treatment of the Apocynaceae of Mexico, the author studied 
the collections of "Echites "of Sesse and Mogino during July 1995 at the Madrid 
herbarium (MA), and from material borrowed from the Field Museum (F) 
during December 1996 in order to locate and observe the types of their 
Echites species. These observations revealed the following novelty. 

The type of the species Echites holosericea Sesse and Mog. has up until now 

Sida 18(1): 237-239. 1998 


SiDA 18(1) 




OB °~ ~i 


ZB " ^ 



C'^ /^^ ^ ^* ^* t*^ J 

/^J-tit^ii* f - ' ^ /£4.,^ZCc/^-<--^- ^l l*^lh^^ 

.1 S,.vso.<i. 0--ini.. vt .MaiJ.'i-ad.. I. .I..V 




NECaii^f MO 



Ji -i ^1 

Fic;. 1 . Holotypc <.->{ Echitcs holosericea Sesse & Mo^. 

Williams, A new combination in Mandevilla 239 

not been formally identified and presented. The typification of £, holosericea 
has been made difficult because there are no specimens so labeled among 
the Sesse and Mogifio collections at F or MA. Many of the posthumously- 
published species names of Sesse and Mogifio are not the same ones that are 
written on the specimen labels (Fuertes & Fryxell 1993). Sesse and Mogino 
(1894) described the species as having opposite subcordate leaves with rusty 
red-white tomentum covering all parts of the plant and white flowers with 
a hairy tube arranged in a compact raceme. Only one specimen (Fig. 1) in 
the Sesse and Mogino collections matches this description. This specimen 
was labeled by them as ''Echites tubulosci N". The name E. tubulosa was never 
validly published by them, however, the letter "N "after the name indi- 
cates Sesse and Mogino considered this specimen to be a new species (Fuertes 
& Fryxell, 1993). Since only one specimen matches the description oiEchites 
holosericea and has a letter "N "on the label it is considered here to be the 
holotype of £. holosericea. 

In 1936, while the Sesse and Mogino collections were on loan to F (McVaugh 
1 980), the specimen disscussed above was identified by Woodson as Mandevilla 
sertuligera Woodson. This identification was later confirmed by the present 
author. With the presentation of the above type it becomes evident that 
the of Sesse and Mogiiio name has priority over Woodson's. A new combi- 
nation is thus required and is accordingly made above. 

The closest relative o'l Mandevilla holosericea is believed to be AI. syrinx 
Woodson. The two species appear to have a close resemblance to one an- 
other. Indeed, there are two sheets of the same collection {Pringle 13106, 
US) that Woodson annotated differently, one sheet he annotated AI. sertuligera 
and the other AI syrinx. Woodson (1938) keyed M. sertuligera apart from 
AI syrinx by its corymbose inflorescence (vs. racemose) and sepals 4—6 mm 
long (vs. 2-3 mm). Both Pringle specimens key more favorably to AI. sertuligera 
(now AI. holosericea). At present I have only observed paratypes of AI, syrinx 
(having been unsuccessful at locating the type at MO), I am inclined, how- 
ever, to recognize the two species as undifferentiated and regard them both 
as a synonym of AI holosericea. 


Fi]i-ini;s, J. and P.A. Fryxell. 1993. Types of names of New World Malvaceae of Lagasca, 

Ortega and Sesse and jVIo^ino. Taxon 42:345—354. 
JMcVauch, R. 1980. Botanical results of the Sesse and Mogifio expedition (1787-1803). 

II. Tlie icones florae iVIexicanae. Contr. Univ. Michigan Herb. 14:99—140. 
SESSfi, M. and J.M. MogiNO 1887. Flora Mexicana. Mexico. 

1894. Flora Mexicana, ed. 2. Mexico. 

Woodson, R.E., Jr. 1938. Apocynaceae. In: N.L. Britton et al., eds. North American flora. 

New York Botanical Garden, New York. 29:103-192. 

240 SiDA 1S(1) 


Fautin, Daphne Gail, Douglas J. Futuyma, and Frances C. James, eds. 
1997. Annual Review of Ecology and Systematics. Volume 28. 
(ISBN {)-cS243-l427-l, hbk). Annual Reviews Int., 4139 El Camino 
Way, P.O. Box 10139, Palo Alto, CA 94303-0139. $60.00. 712 pp. 

Volume 28 is the lisluiI variety ot papers reviewing the tields oi ecology and systemarics. 
The following list ot 25 articles reflects the diversity ol contributions in this 1997 Annual 
Review. (A)ntents: Molecular population genetics ot social insects. Evolution ol eusociality 
in termites. Evolutionary genetics and genetic variation of haplodiploids and x-linked genes. 
Dissecting global diversity patterns: Examples from the Ordovician Radiation. A com- 
parison of alternative strategies for estimating gene flow from genetic markers. The evolu- 
tion of morphological diversity. Insect mouthparts: Ascertammg the paleobiology of in- 
sect feeding strategies. Haldane's Rule. Echinoderm larve and phylogeny. Preserving the 
information content of species: Genetic diversity, phylogeny, and conservation worth. Theoretical 
and empirical examination of density-de|iendent selection. Toward an integrarion of land- 
scape and lood web ecology: The dynamics of spatially stibsidized food webs. Settlement of 
marine organisms in flow. Species richness of parasite assemblages: Evolution and patterns. 
Hybrid origins of phmt species. Evolutionary genetics of life cycles. Phylogeny estimation 
and hypothesis testing using maximum likelihood. Species ttirnover and the regulation of 
trophic structure, Extinction vulnerability and selectivity: Combining ecological and pa- 
leontological views. Tree-grass interactions in savannas. Plant competition tmdergrotmd, 
IMale and female alternative reproductive behaviors in fishes: A new approach using inter- 
sextial dynamics. The role of hybridizcuion and introgression in the diversification of ani- 
mals. The ecology of interfaces: Riparian zones. Allometry for sexual size dimorphism: Pattern 
and process in the coevolution of body size in males and females. 

The usLial indexes follow: Subject Intlex, Cumulative Index of Contribtiting Atithors, 
and a Cumulative Index of (Chapter Titles, Volimies 24-2H. 

Janick, Jules, ed. 1997. Plant Breeding Reviews. Volume I 5. (ISBN 0- 
471-18904-9, hbk). John Wiley & Sons, Inc., 605 Third Avenue, 
New York, NY 10158-0012. $145.00. 395 pp. 

This volume of Plant [breeding Reviews is eletlicatetl to Arnel R. Ilallauer, scientist, maize 
breeder, .uul c]uantitative geneticist. The contents start with a dei-lication to Dr. Hallauer: 
1 ) Detlit at ion: Arnel 11. Hallatier, Scientist, Maize Breeder, Quantitative geneticist. The 
remaining eight chapters cover a variety ol areas in plant breeding. Contents: 2) Molecules 
involved in self-incompatibility in flowering plants. 3) Genetic mosaics and plant improvement. 
4) Quaiitirative trait loci: Separating, pyramiding, antl cloning. 5) Doubleil haploid breeding 
in cereals. 6) Spelt: Agronomy, genetics, and breeding. 7) Cowpea breeding. 8) Recurrent 
selection in soybean. 9) Gene action and plant breeding. All of this followed by a Stibject 
Index, Ctmiulative Subject Index and a Cumulative Contributor Index for volumes 1-15. 

Sii:)A 18(1): 240. 1998 





Pure-Seed Testing, Inc. 

P.O. Box 449 

Hubbard, OR 97032, U.S.A. 

joseph@turf-seed. com 


Herbarium (BRCH) 

Botanical Research Center 

R 0. Box 67 17 

Bryan. TX 77805-6717. U.S.A. 

sdjones@startel. net 


Trachypogon mayaensis, a new species from the Maya Mountains in Belize, is described 
and illustrated. A key is provided to separate it from other species in Central America. 
Trachypogo?! mayaernis is readily distinguished from other Central American species by its 
annual life cycle, height, leaf blade length and width, and prominent and conspicuous sheath 
auricles/ligules, that can reach to 6 cm in length. 


Se describe y ilustra una especie nueva, Trachypogoii mayaensis., de las montaiias Maya de 
Belize. Se presenta una clave para separarla de las otras especies Centro-americanas. T. mayaensis 
se distingue facilmente de ortras especies Centro-americanas por su ciclo de vida anual, 
altura, longitud y anchura de las laminas foliares, sus Ifgulas con aurfculas prominentes y 
cnnspicuas, y que pueden Uegar hasta 6 cm de largo. 

Trachypogon, a genus of approximately six species, is found in tropical and 
subtropical America and Africa (Davila 1994). Two species are found in Central 
America: T. spicatus (C. von Linne) K.E.O. Kuntze and T. vestitus N. Andersson. 
Trachypogon spicatus [syns. = T. montufari (K. Kunth) C. Nees von Esenbeck; 
Trachypogon palmeri Nash; T . plumosus (F. von Humboldt & A. Bonpland ex 
C. von WiUdenow) C. Nees von Esenbeck; and T. secimdus (J. Presl) F. Lamson- 
Scribner} is found from southern Texas in the United States to Argentina, 
and in Africa. Trachypogon vestitus is found from Honduras to Brazil. A col- 
lection o'^ Trachypogon was made in the Maya Mountains in Belize that could 
not be assigned to any known species and is here described as new. 

SiDA 18(1): 241-245. 1998 

242 SiDA 18(1) 

Trachypogon mayaensis Wipff & S.D.Jones, sp. nov. (Fig. 1). 'r> pus: BELIZE. 
Cayo District: 1 .7 mi SE from the c-nrrancc gate inside Mountain F^ine Ridge (SSW 
of Belmopan) in the JVIaya Mottntains, disturbed moimtain pine forest, associates: 
Pitii/s, Andropo^i^on, Paniciim, Axonopiis, Sderici, D'uid'ici, and Aristula\ occasional, L9 Jan 
1991, S.D.Jones 10489 & R. Oliver (holotype: MICH; isotypps: BRCH, MC), US). 

Ciramen anntitun caespitostmi, culmis ( I .59— )1 .95—2.10 m altis; auriculis (3— )4— 5(— 6) 
cm longis (atirictilae ligulae adnatae); tolia lamiiiis (sC— )43-''il cm longis; inflorescentia 
9—30 cm longa. 

Plants (1.59— )1. 95— 2.10 ni rail, robust annual, densely cespitose, erect. 
heaves cauline (measurements taken from mid-culm leaves); nodes ca. II, 
appressed pubescent with trichomcs to 2 mm long, rooting at lower nodes; 
sheaths with ascending to appressed scattered trichomes or glabrous, when 
pubescent, trichomes 1.3—5.5 mm long, becoming denser at apex of sheath; 
collar glabrous; sheath auricles (3— )4— 5(— 6) cm long, erect and adnate to the 
ligules, appressed pubescent or glabrous; ligules (3— )4-5(-6) cm long, mem- 
branous, firm, brown, veined, appressed pubescent attenuate, adnate to the 
sheath auricles; blades (30— )43— 61 cm long, 6—7.5 mm wide (measurements 
taken from mid-culm leaves), flat, apically long attenuate and basally long 
cuneate with the basal portion becoming involute, antrorsely scaberulous 
on both surfaces, margins antrorsely scabrous. Inflorescence 9—30 cm long, a 
spicate raceme or a panicle with two racemose branches; central axis of spi- 
cate raceme or the racemose branches (if a panicle) short pubescent, intern- 
odes 3—4 rnm long (in middle); one pedicellate spikelet terminating spi- 
cate raceme or branch. Spikelets paired, one (lower spikelet) of each pair short 
pedicellate, persistent, awnless, staminate, dorsally compressed; other spikelet 
(upper sj:»ikelet) of pair, longer-pedicellate, deciduous, perfect-flowered, awned, 
nearly terete. Florets without paleas. LOWER (SHORT PEDICELLED) 
SPIKELETS: staminate, sometimes with a vestigial ovary present; the lower 
spikelet either absent or rudimentary in the lowest 4—5 pairs of spikelets in 
the inflorescence or racemose branch. Pedicels 1—2 mm long, pubescent. Spikelets 
6.5-7.6 mm long, 1 — 1.6 mm wide, pubescent] first glumes 6.5-7.6 mm long, 
1—1.6 mm wide, 9— 11 -veined, coriaceous, partially enclosing rest of spike- 
let, narrowly elliptic, sparsely to densely short pubescent, keeled, the keels 
strigose-ciliate, apex bifid, the teeth 0.2—0.3 mm long; second glumes 6.5— 
7.6 mm long, ca. 1.4 mm wide, 3-veined, narrowly elliptic, the margins 
overlapping, ciliate; Lower Floret: lenmias 5.2—5.5 mm long; O-veined, hya- 
line, ciliate on margins; /w/(?i:?j absent. Upper Floret: lemmas 4.7—4.9 mm long; 
3-veined, hyaline, ciliate on upper margins; paleas absenr. UPPER (LONG 
PEDICELLED) SPIKELETS: perfect flowered, awned. Pedicels 2.5-3.1 mm 
long, pubescent. Spikelets (including callus) 9-5—10.6 mm long, 1-1.4 mm 
wide, pubescent; callus., at base of spikelet, 1.8—2.0 mm long, pubescent 
with trichomes to 3.0 mm \ong\ first glumes 7.5—8.5 mm long, 1—1.4 wide, 

WiPl-F AND Jones, A new species of Trachypoyon from Belize 


Fic. 1. Trachypogon mayaensis [S.D.Jones !04i^9 & R. Oliver (BRCH)}. A. Habir (bar equals 
5 cm). B. Section of culm showing the auricle/ligule (bar equals 1 cm). 

244 SiDA 18(1) 

9— 11-veined, coriaceous, margins involute and partially enclosing rest of 
spikelet, elliptic, densely short pubescent throughout, lateral veins near apex 
strigulose, apex rounded and ciliolate; second glumes 7.7-8.9 mm long, 0.9— 
1.3 rnm wide, 3-veined, coriaceous, narrowly elliptic, sparsely pubescent 
between veins; upper margins overlapping and ciliate, apex broadly acute 
and ciliate; Lower Floret: neuter (i.e. without reproductive structures); lem- 
mas 6.5—7.3 nim long, 1.1—1.2 mm wide, 2-veined, hyaline, ciliate on up- 
per margins, apex truncate and c\\\?iX.t\ paleas absent. Upper Floret: perfect; 
lemmas 6—6.9 tnm long, 0.7—1.0 mm wide; glabrous, the lower 1/3 is 3- 
veined, hyaline and easily tearing, the upper 2/3 is 0-veined, subcoriaceous 
to coriaceous and flattened, turning into a terete awn; awn 38—50.5 mm 
long, twice geniculate, pubescent from base to second h^t^c^di; paleas absent. 
Stamens 3, anthers ca. 3.8 mm long, 1-1.1 mm wide. Chromosome number 

Etymology. — The specific epithet refers to the Maya Mountain Range in Belize. 

Phenology. — November-February. 

Distribution. — Known from the Maya Mountains in the Cayo District of 
Belize and from the state of Oaxaca in Mexico. 

Additional specimens examined: BELIZE. Cayo District: Rio Privacion, JMountain Pine 
Ridge, 26 Feb 193 1 , H.R Bartktt 1 1 785 (MICH). MEXICO. Oaxaca: District of Tuxtepe; 
Chiltepec and vicinity, alt. ca. 200 m, 15 Nov 1941, G. Martinez Calderon 812 (MICH). 


(Modified from Davila 1994) 

1. Plants robust annuals, 1.6—2.10 m tall; auricies/liguies (3— )4— 6 cm long; 

leaf blades (30— )43— 61 cm long, 6—7.5 mm wide T. mayaensis 

1. Plants perennial, less than 1.5 m tall; auricles/liguies 2(-5)cm long or less; 

leal blades less than 30 cm long, 1—5 mm wide 2 

2. Plants glabrous or rarely with basal sheath sparsely pubescent; inflores- 
cence a spicate raceme, rarely a panicle with two racemose branches T. spicatus 

2. Plants with sheaths and blades conspicuously pubescent; inflorescence a 
panicle with 3 (rarely 2) racemose branches or rarely a spicate raceme 
T. vestitus 

The closest relative of Trachypogon mayaensis is probably T . spicatus., from 
which it differs by the characters given in the above key; Trachypogon palmeri 
Nash ( = 'r. spicatus) is the name given to longer liguled forms of T. spicatus 
in Mexico. There is also a taxon in Brazil with long ligules, T. macroglossus 
Trinius, but this taxon is perennial with very narrow, involute leaf blades. 

Seeds of T. mayaensis were planted in the greenhouse, of the five plants to 
germinate, only one survived the transplanting to a larger container. Once 
the plant flowered, it began to branch at the lower aerial nodes (typical of 
annuals), but eventually the entire plant died and there were no new tillers 
produced from rootstock. This was also observed in the field. The original 

WiPFF AND Jones, A new species of Trachypogon from Belize 245 

material was collected in November 1994, but when we returned to the 
same site in June 1995, there was no sign of this taxon, though the other 
associated perennial grasses were still present. 


We thank Paul A. Fryxell (TEX) and Tony A. Reznicek (MICH) for their 
assistance with the Latin diagnosis and review of the manuscript; Paul A. 
Fryxell (TEX) for providing the Spanish translation of the abstract and Tony 
A. Reznicek (iVIICH) for loaning specimens oi Trachypogon; and Guy L. Nesom 
(SHST) thank for his initial review of the Latin description. We would also 
like to thank Kelly Allred (NMSU) Gerrit Davidse (MO), WE. Fox, III 
(formerly at TAES), Gretchen D.Jones (USDA, AWPMRU) and Paul Peterson 
(US) for their review and suggestions. We would like to acknowledge the 
generosity of the late Royce L. Oliver (BRIT) for making the collecting trip 
to Belize possible; his generosity is greatly appreciated. We would also like 
to acknowledge Ben Shaw for the illustration. 


DAvii.A, P. 1994. Trachypogon Nees. In. G. Davidsc, M. SousaS., and A.O. Chacer, etls. Flora 
Mesoamericana, Vol. 6, Alismataceae a Cyperaccac. Universidad National Autonoma de 
Mexico, Mexico City, Mexico. Pp. 3iSO-381. 

246 Si DA 1H(1) 


Holm, Lhroy, Ji.rry Doll, Eric Holm, Juan Pancho, and James Hbrberger. 
1997. World Weeds: Natural Histories and Distribution. (ISBN 
0-471-04701-5, cloth). John Wiley & Sons, Inc., 605 Third Avenue, 
New York, NY 10158-0012, U.S.A. $195.00. 1 129 pp. Illustrated. 

Tlie LiLitliors have jiresenced most ot rhc known biology of each species, maps oi their dis- 
tribtition in more than 100 cotmtries, illustrations designed to lielp witli identiHcation, 
and an index of common names, and a bibli()gra|-ihy of 3300 references to lead students 
and researcliers ro further details that are contamed in the jTrmcipal papers. From the cover: 
"The culmination of four decades of gloloal research, World Weeds jiresents comprehen- 
sive and Lip-to-date information on over 100 weeds-addressing recent changes in such ar- 
eas as cro|T tillage methods, herbicide Lise, and agricultural runoff. This monumental work, 
featLiring a wealth of original data from the authors, provides extensive coverage of the 
know biology of each species. Each entry contains a full botanical description, plus impor- 
tant details on iiabitat recjuiremenrs and distrif^ution, seed production, ecology, physiol- 
ogy, croji impact, and more." It is generously supplied with excellent line drawings. 

Hodgson, Geoifrey M. 1995. Economics and Biology. (The International 
Library ol Critical Writings in Economics). (ISBN 1-85898-050-X, 
hbk). Edward Elgar Publishing Limited, England and Edward Elgar 
Company, Old Post Road, Brookfield, VT 05036, LJ.S.A. $189-95. 
606 pp. 

The International Library of Critical Writings in Economics series is an essential reference 
soLUce for students, researchers and lecturers in economics. Economics and Biology is number 
50 in the series. There are six parts each divided into several chapters. Part 1: Biological 
and Mechanical Analogies. Part II: Economics and Sociobiology. Part III: Classical Eco- 
nomics aiul the Darwinian Revolution. Part IV: Alfred Marshall and Economic Biology. 
Part V: Evolution, Optimization antl Rationality, j-'art Vi: Biology and Modern Econom- 
ics. A name index follows. 

SiDA lcS(l): 246. 1998 



The Nat//re Conservancy 

P.O. Box 2267 

Chal)dWll, NC 27313-2267, U.S.A. 


Department of Botany 
National Museum of Natural History 

Smithsonian Institution 
Washington, DC 203(^0-0166, U.S.A. 


The Sporoboliisfloridamn complex is defined to include five North American species. Keys, 
descriptions, distributions, illustrations, and habitat information are provided for: S. curtissii, 
S. floridaniis. S. silveanus. S. teretifolius. and a new member of the complex from North Caro- 
lina, South Carolina, and eastern Georgia, Sporoholiis pinetoruvi Weakley and P. M. Peterson. 
A lectotype is chosen fori', jlrnidantn. The taxonomic and ecological relationships ol these 
five species, as well as the related S. juiiceus and S, beterokpis, are compared and discussed. 
Spomlmliii cnrtisiii, S. floruUinii.s. S. pinetorum, S. silveanus, and S. teretifolius are all relatively 
narrow endemics of various portions of the southeastern Coastal Plain. Each ot these five 
species is the locally dominant or codominant grass in fire-maintained pinelands with open 
canopies q{ Pirius palustris, P. serotina. and/or i^ elliottii y3.r:. elliottii. The geographic distri- 
butions and many ecological requirements of the 5. curtissii. S. floridarius. S. pinetor//ni, and 
S. teretifolius overlap, but they can be separated on a hydrologic gradient. 


El complejo Sporoholus floridanus es definido para incluir cinco especies de Norte America. 
Se proporcionan claves, dcscripciones, distribucion, ilustraciones, e inlormacion del iiabitat 
para: S. curtissii, S. floridanus, S. silveanus, S. teretifolius, y un nuevo miembro del complejo 
de Carolina del Norte, Carolina del Sur y del este de Georgia, Sporobolus pinetorum Weakley 
& P.M. Peterson. Un lectotipo es escogido pzitd^S. floridanus. Eas relaciones taxoncjmicas y 
ecologicas de estas cinco esjiecies, asi como de las especies alines S. junceus j S. heterolepis, 
son comparadas y discutidas, Sporobolus curtissii, S. floridanus. S. pinetorum. S. silveanus, y S. 
teretifolius son todas relativamente endemicas restringidas a varias partes del sureste de la 
llanura costera. C^ada una de las cinco especies, es el pasto localmente dominante o codominate 
en terrenos de i^inos mantenidcjs por quemas con dosel abierto de Pinus palustris, P. serotina, 
y/o P. elliottii var. elliottii. La distribucion geografica y muchos de los requerimientos ecologicos 
de S. curtissii, S. floridanus, S. pinetorum, y S. teretifolius se solapan, pero pueden ser separados 
en base a un gradiente hidrologico. 

Sporobolus R. Br. is a worldwide genus of approximately 160 species oc- 
curring in the tropics, subtropics, and warm temperate regions (Clayton & 

SiDA 18(1): 247-270. 1998 

248 SiDA 18(1) 

Renvoize I 986). In the New World the genus is represented by approxi- 
mately 45 species that generally occur on disturbed habitats, i.e., roadside 
to open prairies and savannas (Peterson et al. 1995, 1997; Peterson et al., 
in press). Sporohoh/s is characterized by having spikelets with one floret, 1- 
veined lemmas, fruits with free pericarps, and ligules with a line of hairs. 
These characteristics also are found in two other genera, Calamovilfa (A. 
Gray) Hack, and Crypsis W.T. Alton. These three genera seem to share a 
common ancestor and are the only New World members included in the 
subtribe Sporobolinae (Chloridoideae; Eragrostideae). 

The species oiSporobolus occurring in the southeastern United States form 
a heterogeneous assemblage that may be informally divided into groups based 
on characteristics of the inflorescence, spikelet, plant longevity, and gen- 
eral aspect. One of these groups, characterized as long-lived, perennial, clump- 
forming species, with open panicle inflorescences and relatively large spikelets, 
consists oiS.floriclani/s Chapman, S. airthsit (Vasey ex Beal) Small ex Scribner, 
S. teretijolius Harper. S. silveanus Swallen, S. pinetorum Weakley & P.M. Peterson 
(a new species described here), and, more peripherally, S. junceus (P. Beauvois) 
Kunth and S. heterolepis (A. Gray) A. Gray. In the course of conducting con- 
servation, taxonomic, ecological, and herbarium studies in the southeast- 
ern United States, it has become apparent that this group is poorly under- 
stood and has been much confused by earlier authors, current collectors, 
and field workers. As a consequence, an overlooked species has remained 
unnamed. We will try to clarify this situation with a detailed discussion of 
the five members of what we call the S. flortdaniis complex {S. flondanus, S. 
curttssit. S. teretijoln/s. S. silvean//s, and S. pinetoriini), with less detailed dis- 
cussion of .S. junceus and S. heterolepis. 


Because of the general similarities of the taxa in the Sporobolus jloruhvius 
complex, there has been widespread confusion about their circumscription 
and distribution, and much erroneous information can be found in earlier 
systematic treatments and in the ecological literature, and numerous 
herbarium specimens are misidentified. A review of previous treatments of 
the complex and its closest relatives will help define problems resolved in 
this paper. 

The two morphologically peripheral taxa were the first to be named. 
The species currently known as S. junceus was described by Beauvois in 1812 
in Heleochloa, and transferred to Sporobolus by Kunth in 1 829- Sporobolus heterolepis 
was first described as Vilfa heterolepis by A. Gray in 1835, and transferred 
by Gray to Sporobolus in 1848. 

Weakley and Peterson, Sporobolus floridanus complex 249 

The first of the core members of the S. floridanus complex was named by 
Chapman (I860) as S. floridanus. It is fairly well characterized in his de- 

"S. floridanus, n. sp. Panicle clilhise, large; spikclets (purplish) on long hair-like stalks; 
glumes acute, the lower one barely shorter than the obtuse paleae, the upper a third longer; 
leaves rather rigid, flat, pungent, very rough on the edges— Low pine barrens. Middle and 
West Florida. September.— Ctilm 2—4 feet high. Leaves 1—2 feet long. Panicle 1 — 1.5 feet 
long" (Chapman I860, 1883, 1897). 

The description, the location in Florida, and specimens collected by Chapman 
at the time make the identity of ^. floridanus clear. The only species occur- 
ring in Florida likely to be confused with S . floridanus would be S. curtissii, 
but it is contradicated by various parts of Chapman's description, especially 
the "panicle ... large," the markedly unequal glumes, the rough-edged leaves, 
and the length of the blades and the panicle. No type specimen was cited, 
however, and we lectotypify S. floridanus below. 

The next to be named was S. curtissit, as S. floridanus var. curtissii (Beal 
1896). A year later, Lamson-Scribner elevated the taxon to specific rank (Lamson- 
Scribner 1897). The distribution of 5". floridanus was given as "moist pine 
barrens near the coast, North Carolina to western Florida," apparently on 
the basis of G. McCarthy's specimen at US (cited below) from Wilmington, 
North Carolina, which is actually S. pinetorum. We know of no specimens of 
S. floridanus from locations north of southern South Carolina. 

Harper (1901) made a characteristically idiosyncratic addition to infor- 
mation on the group with his collection of i', floridanus in Sumter Co., Georgia, 
about which he commented "not definitely known outside of Florida be- 
fore." He elaborated on the toughness of the leaves mentioned by Chapman 
and stated "I made some tests of their strength. A leaf from the first collec- 
tion (no. 547) 5.5 mm. wide, not twisted, and perfectly dry, sustained a 
weight of 27 pounds without breaking." 

In 1906, Harper described S. teretifolius from collections in Georgia, com- 
menting that it is "a frequent and characteristic inhabitant of moist pine- 
barrens in the Altamaha Grit region," and that "it is unmistakable when 
seen in the field" (Harper 1906). He provides excellent and clear character 
differences between S. teretifolius, S. curtissii, and S. floridanus, as well as an 
excellent illustration of the highly distinctive leaf cross-section. 

The first two editions of J.K. Small's flora (1903, 1913) treated S. curtissii 
and S. floridanus, and gave generally accurate information about morphol- 
ogy, habitat, and distribution. Sporobolus floridanus was stated as occurring 
in "Georgia and northern Florida," and S. curtissii was found in "Georgia, 
Florida, and Alabama." Small then added S. teretifolius (Small 1933). The 
habitat of all three species was given as "moist pinelands, Coastal Plain," 

250 SiDA 18(1) 

with distributions of "Fla. and Ga" for S. floridanus, "Fla. to N.C." for S. 
c//rtissii, and "Cja." for ^. teretifoliiis. The extension of the distril:)ution of 5". 
airtissii to North Carolina was apparently based on North (Carolina speci- 
mens of S. p'Dietor/oii misidentified as S. curt'nsii. In his key, Small (1933) 
distinguished the three species as follows: 

Leaf-blades narrowly involute S. ttretifali/is 

Leaf blades flat. 

Pedicels appresscd to the branches: leaf-blades al^toiit I mm wide S. c/irtisiti 

Pedicels spreading: leal-blades 2-4 mm wide S. floridanin 

Although this key generally serves to distinguish the three species, it has 
been, in part, responsible for continued taxonomic confusion about the three 
species and a fourth, 5", ptnetornni. The blades of 5, teretijolii/s are anatomi- 
cally oval, not involute; the blades of .V. cnrtissii, S. floruhiniis, and .V. phietomvi 
are flat, becoming involute when dry, either during drought conditions in 
the field, or as a result of drying for herbarium specimens. The narrow blades 
of S. airtmii and S. pmetorum usually fold when dried in a plant press. Specimens 
of .V. curtissii and S. pimtonmi often have involute blades that superficially 
resemble those of 5. teretifolius. A second problem with the key involves the 
width of the blade; both S. c/irtissii and S. pmetorum have blades regularly 
reaching and sometimes exceeding 2 mm in width (when flat), and S. floriclaii/ts 
characteristically has a leaf blade 3-10 mm wide (though the very narrow- 
est blade on a plant may be as narrow as 2 mm wide). Therefore, specimens 
of S. cnrtissii and the heretofore unnamed S. pmetorum have often been in- 
correctly identified as S. teretifolins (by taking the first lead) or S. floridanus 
(by correctly taking the second lead in the first couplet, but then incor- 
rectly choosing the second lead in the second couplet by placing too much 
emphasis on a blade width of ca. 2 mm). 

Hitchcock (1935) treated S. fhyridanm ("low pine barrens, Georgia and 
Floricfa"), S. cnrtns'i'i ("dry pine barrens. North Carolina, Georgia, and Florida"), 
and S. teretifoliz/s ("moist pine barrens, Georgia"). Once again, the inclusion 
of North Carohna in the distribution of .S', cnrtissii was apparently based on 
miss identification of specimens of S. pmetoriini. 

Blomquist's ( 1 9 IH) book on the grasses of North Carolina treated all material 
of this complex in North Carolina as S. a/rtissii, but states "According to 
Swallen (1941) the North Carolina plants assigned to this species may be- 
long to S. floridcinns Chapm." As will be seen below, neither species has been 
documented for North Carolina; instead. North Carolina is within the dis- 
tribution o{S. piiietor/nii (undescribed at the time and in some ways gener- 
ally intermediate in characteristics) and S. teretijolms {not collected in North 
Carolina until 1 991). 

Swallen (1941) added S. silveciuns to the group, based on material from 
eastern Texas. Its distribution in Texas and the western (nilf Coastal Plain 

Weakley and Peterson, Sporobolus floridanus complex 251 

of Louisiana makes it allopatric relative to other members of the S. floridanus 
complex. Swallen contrasted it with S. floridanus and S. teretifolius, yet vari- 
ous characters (such as the appressed and larger spikelets) suggest a closer 
relationship to S. curtissii. It also seems to show a clearer connection to 5". 
heterolepis than do other members of the 5^, floridanus complex, a relation- 
ship which seems especially plausible given its proximity to S. heterolepis 
and its habitat preferences for barrens with prairie affinities. 

Hitchcock and Chase (1950) provided the only treatment to date with 
as many as four of the species in the complex: S. floridanus ("low pine bar- 
rens, North Carolina to Florida"), S. curtissii ("dry pine barrens. North Carolina 
to Florida"), S. teretifalius ("moist pine barrens, North Carolina and Geor- 
gia"), and S. silveanus ("open woods, western Louisiana and eastern Texas"). 
The attribution of 5^. floridanus, S. curtissii, and S. teretifllius to North Caro- 
lina are all based on misidentifications oiS. pimtorum. This is because of the 
generally intermediate morphology oiS. pinetorum, and because the key was 
not constructed very carefully for even the then-known members of the group. 
Among the problems are that S. silveanus (which has markedly appressed 
spikelets) can only be reached by following the lead for "spikelets not ap- 
pressed, the branches and pedicels somewhat spreading"; S. floridanus is separated 
from S. silveanus by having glumes "about equal" rather than "unequal," 
when they actually have similarly subequal first glume to second glume 
ratios; S. teretifllius is separated from S. floridanus by the accurate but often 
mis-interpreted "blades terete vs. blades flat or folded" character; and S. 
curtissii is separated from the others by a subjective and difficult to inter- 
pret couplet about pedicel length and orientation. 

Radford et al. (1964), by contrast, recognized only one species {S. teretifldius) 
in the complex as occurring in North Carolina and South Carolina. Unfor- 
tunately, all material seen by them was actually the undescribed S. pinetorum. 
Radford et al. (1968) added S. floridanus (alleged to occur in pinelands in 
Lancaster County, South Carolina) and attributed S. teretifllius to "savannahs" 
in thirteen counties in southern North Carolina and northern South Carolina. 
Their key distinguished S. teretifllius from S. floridanus by "blades terete or 
subterete" vs. "blades flat or folded." All material seen by them was actually 
5, pinetorum, often with narrow folded blades so as to superficially mimic S. 
teretifldius. Additionally, many of the county records are based on misidentified 
specimens oiCalamovifla brevipilis (Torrey) Scribner, and likewise specimens 
supposedly documenting county record distributions for Calamovifla brevipilis 
are in many cases actually Sporobolus pinetorum. In truth, both Sporobolus pinetorum 
and Calamovilfl brevipilis occur (or formerly occurred) in most or all counties 
in southeastern North Carolina and in the adjacent counties of South Carolina. 

Godfrey and Wooten's (1979) manual of southeastern United States wetland 
plants treated only S. floridanus and S. curtissii, inexplicably omitting S. teretifalius 

252 SioA 18(1) 

(which occupies wercer habitats than S. ani/s.w/) and S. nlveanus (which oc- 
curs in wetland situations in Louisiana, part of the geographic range of their 
flora). Because of this, their treatment is of limited value in understanding 
this group. 

Brown (1993) addressed confusion between S. silveaniis, the most west- 
ern species in the complex, and S. heterolepis, resulting in the deletion of ^. 
heterolepis from the floras of Texas and Louisiana, and the addition of 5. silveanm 
to the flora of Oklahoma. 


1. Panicle branches tlisrinctly whorled ac lower nodes S. junceus 

1. Panicle branches single at lower nodes (though a Few branches may be ir- 
regularly approximate in pairs or threes) 2 

2. hirst ghime scaberulous, subulate above an expanded base; spikeiets gray 
to nearly black; base of |ilant relatively fibrous; grain spherical; plants of 
rocky barrens and prairies of physiographic provinces inland from the Coastal 

Plain S. heterolepis 

2. First ghime glabrous, lincar-lanceohxte to lanceolate, the base not abruptly 
expainled; spikeiets purplish (fadmg tan); base of plant smooth and hard, 
made up of the indurated leaf bases; grain oblong (when presenr, usually 
abortive); plants primarily of pine savannas and seeps of the Coastal Plain 
of North Carolina, South Carolina, Georgia, Florida, Alabama, Louisiana, 
and Texas, bur exrending ftuTlier inland in southern Oklahoma and 

eastern Texas 3 

3. Blades terete or subterete (wiry), 0,5-1.2 mm wide, oval in cross- 
section (dee|-)er than wide), sometimes irregularly channeled for por- 
tions of their lengths (but lacking any flat portion), margins smooth, 
distal portions often curling and twisted; pedicels witii scattered as- 
cending hairs 5. S. teretifolius 

3. Blades flat, O.S-IO mm wide, flat or V-shaped in cross-section (much 
wider than deep), with free margins their entire length, margins scaberulous 
(glabrous to scaberulous in S. ciirtisui), disral portions normally stiff 
and straighr (note that the blades of the narrower-leaved species can 
appear superficially wiry); pedicels without scattered ascending hairs, 

either glabrous to scaLoerulous or scabrous 4 

4. Lower ghime usually as long or longer than the upper glume with 
lower/upper ratio averaging 0.90-1 . i 5; culms 30-80(-90) cm tall; 
panicle 1 0-25 cm long; pedicels 0.5— K-H) mm long, appre.ssed, usually 
shorter than the spikeler; leaves less than 30 cm long, smooth on 

the margins 1. S. curtissii 

4. Lower glume tisually shorter than the upper glume with a lower/tipper 
ratio averaging 0.60-0.90; c ulms (30-) i5-2(){)(-250) cm tall; panicle 
15-50 cm long; pedicels 1-11 mm long, spreading or appressed; 

leaves mostly more rhan 30 cm long, scaberulous on the margins 5 

5 . Pedicels appressed; lemmas 4.4-6.5 mm long, purple; anrhers 3.5- 
5 mm long; blades 1-2.5 mm wide and bluish-green; jilants known 
west of the Mississippi River 4. S. silveanus 

Weakley and Peterson, Sporobolus floridanus complex 253 

5 . Pedicels spreading; lemmas 3—4.3 mm long, purplish-brown; anthers 
2-3.4 mm long; blades either wider than 2.5 mm, or dark green; 

plants from east of the Mississippi River 6 

6. Blades (2-)3-10 mm wide, bluish-green; panicles (18-)30-50 
cm long, 4-15 cm wide; lower/upper glume length ratio aver- 
aging 0.75-0.90; plants from southctn South Carolina, Geor- 
gia, eastern Alabama, and northwestern Florida 2. S. floridanus 

6. Blades L2-2(-3) mm wide, dark green; panicles 1 5-30 cm long, 
2-6 cm wide; lower/upper glume length ratio averaging 0.60- 
0.80; plants from eastern North Carolina, northern South Carolina, 
and eastern Georgia 3. S. pinetorum 

1. Sporobolus curtissii (Vasey ex Beal) Small ex Scribn., U.S.D.A. Div. 
Agrostol. Bull. 7:142. 1897. (Figs. 1, j; 2). Sporobolus flonclanus wav. iurtissii 
Vasey ex Beal, Grass. N. Amer. 2:290. 1896. Type: U.S.A. Florida: 1883, Curtiss 
S.71. (holotype: MSC; lsotype: US-556876!). 

Caespitose perennials. Culms 30-8()(-90) cm tall, erect, nodes all basal; 
base diameter 1-2 mm, flattened; inrernodes glabrous. Sheaths glabrous to 
appressed hairy, hairs up to 4 mm long; base shiny and endurated; margins 
hyaline; summit with a tuft of hairs, hairs up to 4 mm long. Ligules 0.2- 
0.6 mm long, a line of hairs. Blades 5-22(-28) cm long, 0.8-2(-2.2) mm 
wide, flat to folded or involute, green, remaining green well into winter, 
mostly glabrous above and below; base densely pilose on upper surface, white 
to grayish hairs up to 4 mm long; margins glabrous to scaberulous. Panicles 
10-25 cm long, 2-10(-13) cm wide, mostly open, contracted when imma- 
ture, pyramidal to ovate; main axis glabrous to scaberulous; pulvini in axils 
of primary branches glabrous or occasionally hairy; primary branches 2- 
9(-10) cm long, ascending to spreading 10-80" from culm axis, not florif- 
erous on lower 1/3; secondary branches mostly appressed; pedicels 0.5-4(- 
8) mm long, usually shorter than spikelet, appressed, glabrous. Spikelets 
3.5-6(-6.6) mm long, purphsh-brown. Glumes (2.9-)3.5-6(-6.6) mm long, 
linear-lanceolate, membranous, 1 -veined, equal to subequal; ratio of lower/ 
upper glume length 0.90-1 . 1 5(- 1 .33); lower (2.9-)3.5-6.2 mm long, apex 
acuminate; upper 3-2-6.6 mm long, apex acuminate. Lemmas 3.4—4.5 mm 
long, ovate to lanceolate, membranous, 1 -veined, glabrous; apex acute. Paleas 
3.4-4.5 mm long, ovate, membranous, glabrous; apex acute. Stamens 3, 
anthers 1.5-2.8 mm long, yellow to purplish. Grains 1.1-1.4 mm long, 
fusiform, reddish-brown. 

Common name. — Curtiss' dropseed. 

Distribution and habitat. — Eastern South Carolina south to central pen- 
insular Florida, west to Florida Panhandle. Mesic to dry-mesic pine wood- 
lands, in soils seasonally saturated at the surface or rather well-drained throughout 
the year, usually under Pinus paiiistris, and sometimes also with Quercus spp.\ 
0-100 m. Sporobolus curtissii characteristically occurs in the following Na- 


Si DA 1<S(1) 


Fi(,. I. A-E. Sjmrohfil//s pn/ctoiy/iii ih-d'^.ijd on \\'t:cikU'y s.n., 31 Ali^' 1993). A. Habic. B. Lii^'iile. 
C. Spikelcc. 15. Pistil and stamens. E, Blade, lateral view. F. Sporobohis teyetifoliiis {Wedkky & 
Schafcile }■.!/., 16 JliI 1991). F. Blade, transverse section. G. Sporoho/us s//reai/us {Waller & 
Bci/iml M 28). G. Four spikelets on a branch of the inflorescence. H, I. Sporobolin flor'nhinin 
(Ciirtiis 4034}. H. .Spikelet. I. Blade, abaxial view. J. Sporohoh/s a/rtissii {Diniuiu 785 5). J. 

Wf.aki.f.y and Peti-.RSON, Sporobolus floridanus complex 


Fig. 2. Geographic clistribLition oiSponbDliis a/rtissii {K), Sporobolus pinetnrum (B), and Sporohol//s 
teret if alius (C). 

tional Vegetation Classification plant associations: Vinm palustrislSerenoa repens- 
Vaccinmm myrsinitesi Anstida beyrkhiana— Sporobolus curtissii Woodland; Finns 
palustrhlQiiercus mcana-(2piera(S stellata/A ristula beyrkhiana-Sporobolus juncem- 
Nolina georgiana Woodland; Vinus palmtrh-iPtnus elliottii var. elliottii) /Sporobolus 
pmetm-um- Aster reticulatus-iSporobolus curtissii) Woodland (Weakley et al. 1998). 
Flowering July to November (or less typically at other seasons if stimu- 
lated by fire). 

Comments. — When fertile, S. curtiss'ii is readily distinguished by the short- 
pediceled, appressed spikelets, and by the large lower glume. Vegetatively, 
S. curtissii is distinctive in its short leaf blades and tendency to have gla- 
brotis leaf margins. Post-fire foliage tends to be narrow, stiff, and erect, while 
unburned plants produce leaves that are shorter, wider, and laxer. It occu- 
pies drier habitats than S. floridanus, S. teretifolins, and S. pinetorum, though 
it can be found in mixed populations with all three species. 

256 SiDA 18(1) 

Representative specimens. FLORIDA. Alachua Co.: Gainesville, Coiiihs i.n. (GH). Baker 
Co.: Okisfee Ex|>erimental I'orest, Okistee, 9 jun 1936, Wtitknis I (US). Bradford Co.: 
(FLAS). Clay Co.: 7 mi NW of Midclleburg, abundant on peaty pond pme savanna, 28 Jul 
1967, McDunid 9M' / (BRIT/VDB, G A). Columbia Co.: Lake City, 1 1-1 9 Jul I 'A^■^,Hash 
22 1 3 (GA, GH, NCU, US). Duval Co.: moist pine barrens near Jacksonville, 6 Au^i; & 13 
Nov 1 89-1, C//r//xv 3!8I (GA, GH, US). Manatee Co.: 1996, Alr/M/7/./;/ 1534 (NCU). Okaloosa 
Co.: (reported in Clewell 1985). Orange Co.: Cirasmere, 23 Sep 1898, C&w/;j & Baker J 1 27 
(US). Polk Co.: 1 996, McMi//a?i I 534, (NCU). Putnam Co.: E of Palatka, open field, 26 
Oct 1940, S//i'i'//s 6737 (US). St. Lucie Co.: dry pme barrens, Fort Pierce, 25 June 1939, 
Silve//s 4429 (DUKE). Taylor Co.: wet pine llatwoods, 1 979, GM/frey 77245 (Ci A). GEORGIA. 
Berrien Co.: Ala]-)a Experimental Range, 2 mi S of Alapa, dominant (reported by Lemon 
1949); Brantley Co.: moist open pinelands 1.5 mi W of Nahunta, 1953, Diniani 17036 
(GH). Bryan Co.: Fort Stewart, 1996, McM/7A/// 1061. 1064. 1065 (NCU). Charlton Co.: 
Camp Cornelia, 1992, Ricka- 932 (GH, US). Clinch Co.: pme barrens, Okifionokee [sic] 
pocket, 8 Oct 1938, liyles 396 (DUKE). Coffee Co.: 5 mi south of Ocmulgce River, 18 
May 195i, Cooley 2780 (US). Cook Co.: 2.7 mi NE of Barney on Ga. 76, 7 Nov 1993, 
Sorrie 7786 (NCU). Dougherty Co.: dry sand, fossil dunes. East Albany, 30 Aug 1947, 
Thorne 61 il (CiA). Echols Co.: longleaf pine-palmetto flatwoods near Ga. 94, 17 Aug 
1967, Clewell 2624 (FSU). Glynn Co.: 1996, McMillan 1082. 1082a (NCU). Liberty Co.: 
Fort Stewart, \996, McMillan 1068. 1074. /07_5 (NCU). LongCo.: \996, McMillan 1048. 
1048a (NCLJ). Lowndes Co.: S of Melrose, 4 Sep 1902, Harjm- 1603 (GI 1, US). Tattnall 
Co.: 1 996, McMillan 1081 (NCLJ). Toombs Co.: 1 3 Dec 1 995, Some & Weakley 7801 (GA). 
Wayne Co.: 1 mile S ofjesup, 1 947, Duncan 7855 {GA, GH, FLAS, US). SOUTH CAROLINA. 
Berkeley Co.: Francis Marion National Forest, 21 Jul 1994, Sorrie 807 5 (NCU); locally 
dominant, growing in dense and obviously quite old tussocks in gummy longleaf/loblolly 
flatwoods W of Tiger Corner, 30 May 1995, McMillan & K/elln/ark 10 H (NCU, USCH), 
24 Jul 1996, McMillan 1 748 (CLEMS, NCU, USCH). 

2. Sporobolus floridanus Chapm., Fl. South. U.S. 550. 1860. (Figs. 1, h, 
i; 3). Typi:: U.S.A. FLORIDA. Franklin Co.: Apalachicola, Chapman .Lti. {L\-x:ny\Y\>\-, 
here designated: NY, bar code, 00127474!; iscilhctotypr: US-998263!). 

Robust caespkose perennials. Culms (40-)100-200(-250) cm tall, erect, 
nodes all basal; base diameter 1.5-6 mm, flattened; internodes glabrous. 
Sheaths glabrous to appressed hairy, hairs up to 5 mm long; base shiny and 
endurated; margins hyaline; summit with a tuft of hairs, hairs up to 6 mm 
long. Ligules 0.2-0.7 mm long, a line of hairs. Blades (l()-)25-50 cm long, 
(2-)3-10 mm wide, flat to folded, pale bluish-green and yellowing in age, 
mostly glabrous above and below; base often sparsely hairy, hairs up to 6 
mm long; margins scaberulous. Panicles (1 8-)30-5() cm long, 4-15 cm wide, 
mostly open, contracted when immature, pyramidal to ovate; main iixis scabrotis; 
pulvini in axils of primary branches hairy or glabrous; primary branches 4- 
15 cm long, ascending to spreading lO-yOr* from culm axis, not floriferous 
on lower 1 /3; secondary branches ascending to spreading; pedicels 2-14 mm 
long, usually longer than spikelet, spreading, scaberulous. Spikelets (3.7) 
4-6 mm long, purplish-brown. Glumes (2.5-)2.8-5.7 mm long, linear- 
lanceolate, membranous, 1-veined, subequal; ratio of lower/upper glume 
length ((),60-)().75-0.90(-0.94); lower 2.5-5. 1 mm long, apex acuminate; 

Weakley and Peterson, Sporobolus floridanus complex 


Fig. 3. Geographic distribution of Sporobolus floricLniifS (A) and Sporohnlns itlvmnin (B). 

Upper 3.7-5.7 mm long, apex acuminate to acute. Lemmas 3-4 mm long, 
ovate to lanceolate, membranous, 1 -veined, glabrous; apex acute. Paleas 3- 
4 mm long, ovate, membranous, glabrous; apex obtuse to truncate, rarely 
minutely bifid. Stamens 3, anthers 2-3.1 mm long, purplish. Grains 1.7- 
2 mm long, fusiform, reddish-brown. 

Common name. — Florida dropseed. 

Distribution and habitat. — Southeastern South Carolina south to north- 
ern peninsular Florida, west to Florida Panhandle. Wet to wet-mesic pine 
woodlands, in soils semi-permanently to seasonally saturated at the surface, 
and even in places where water may pond for weeks, usually under Finns 
elliottii var. elliottii. Pim/s palustris. or Taxodt/nn ascendens Brongn., and also 
in seepage bogs, treeless swales, and depressional wetlands in pineland landscapes; 
0-100 m. Sporobolus floridanus characteristically occurs in the following National 
Vegetation Classification plant associations: Pinus palustris— P inns elliottii var. 
elliottiilCtenium aromaticum-A ristida bey rich iana-Sporobolus floridanus Woodland ; 
Pinus palustris— Pinus elliottii var. elliottiitStyrax americanus var. pulverulentusl 
Sporobolus floridanus Woodland (Weakley et al. 1 998). Flowering July to November 
(or less typically at other seasons if stimulated by fire). 

Comments. — This is a common and conspicuous grass within its range, 

258 Sum 1<S(L) 

and is often a dominant to codominant component of wet pinelands. The 
wide leaves with distinctly bluish cast are distinctive in this species. 

Representative specimens. ALABAMA. Houston Co.: (Ciiinn, pers. tomm). FLORIDA: 
Dixie Co.: Ilatwoods near Hines, 26 Aui,' 1 937, West ct uL HH-n4 (FLAS, US). Duval Co.: 
moist pine barrens, near Jacksonville, Oct I 894, Ci/rtiss ]]7^ (GA, GH, NCU). Franklin 
Co.: pine-turkey oak woodland [likely a wetland mclusion in or near this habitat}, 1 mile 
from brid_t;e over Ochlochonee Bay, 14 Jul 1954, Fonl ]} 24 (US). Gulf C]o.: St. joe, open 
pmeland, 5 Oct 1940, Silveus 649H (US). Levy Co.: Fllzen [or Ellzey], Ilatwoods, infre- 
c]Lieiu, 3 Sep 1898, Coiuhs HIH (US). Liberty Co.: AiicJersun 12503 (FSU). Madison Co.: 
1955, Godfrey ^4004 (DUKF, FSU, GH). Wakulla Co.: St. Marks Refu^^e, Some 7^85 
(NCU). Walton Co.: de Funiak Springs, 1 898, Combs 446 (GH). GEORGIA. Baker Co.: 
Jones Ecological Research Station, Ichauway Plantation, Jack Baker Woods, 17 Aug 1 995, 
AlcM///i/>/. Kjellii/ark. & Dreu 1110 (USCH). Berrien Co.: Forest Experiment Station, 5 
mi NE of Tilton, open pme woods, moist sandy soil, 1943, Biswell & Lemon PL-270 {¥LAS, 
and reported in Lemon 1949). Brooks Co.: wet seepage area in upland pine woods, 1.4 mi 
N of Morven, 6 Sep 1967, Pciirdoth & Fuircloth 4i329 (GA, NCU). Bulloch Co.: , 20 April 
1994, Some 7859 (GA, NCU). Cook Co.: i mi SE of Adel, open pme forest, 27 Jun I 957, 
Pohl 7636 (GH). Dooly Co.: open partially drained cypress pond, on W side of railroad, 
2.7 mi SSE of Unadilla, 31 Jtil 195 i, H,/rper. H/nnphrey. & Diinani I6H39 (GA). Echols 
Co.: open pine-palmetto flatwoods on the E side of the Alapaha River, 3.6 mi S of Mayday, 
21 Oct 1967, fcinrloth 497H (GA, NCU). Lanier Co.: open pine woodland alongside LIS 
221, 2.5 mi WSW of Lakeland, 12 Oct 1967, Pa/rc/oth 4930 (GA, NCU). Long Co.: dis- 
turbed pocosin, 6.3 mi W of 301 on Ga. 26 1 , 6 Oct 1962, Bozemai/ & Radjonl 1964 (NCU). 
Lowndes Co.: open longleal pine forest on level terrain 5 mi E of Valdosta, 12 Oct 1963, 
Duncan 22210 (GA). Sumter Co.: moist pine barrens, 31 Aug 1900, Harper "^47 (GH). 
Tattnall Co.: common in slash pine flatwoods clearing, ca, 2 mi S of Glenville, 1 I Sep 
1 972, Godfrey 72107 (FSU, NCU). SOUTH CAROLINA. Jasper Co.: dominant on mesic 
to wet Pi)ii/s ellioltiilP. palnstris savanna on Rains series sand}' loam with Paxville fine loamy 
inclusions, 28 Jun 1995, McMillan & Kjellmark 1035 (CLEMS, NCU, USCH). 

3- Sporobolus pinetorum Weakley & P.M. Peterson, sp. nov. (Figs. 1, a- 
e; 2). Tyvv.: U.S.A. North Carolina. Cumberland Co.: Fort Bragg Army Base, Macridge 
Danger Zone, ecotonal seepages and moist swales in uj:iland Pnii/s palustris savannas 
Weakley s.n., 31 Aug 1993 (hoi.otypi:: US!; lsotypr: NCU!). 

A S. fluridaui laminis foliorum 1 .2-2(-3) mm latis atrovirentibus, inflorescentisl 5-30 
cm latis, culmis 45-120 cm altis, ratione longitudinus glumarum inferiorum/superiotLim 
0. 60-0. 80, recedit. 

Caespitose perennials. Culms (3()-)45-12()(-18()) cm tall, erect, nodes 
all basal; base diameter 1-3 mm, flattened; internodes glabrous. Sheaths 
glabrous to appressed hairy, hairs up to 4 mm long; base shiny and endurated; 
margins hyaline; summit with a tuft of hairs, hairs up to 4 mm long. Ligules 
0.2-0.6 mm long, a line of hairs. Blades 20-50 cm long, 1.2-2(-3) mm 
wide, flat to folded or involute, dark green, remaining green well into win- 
ter, mostly glabrous above and below; base often sparsely hairy, hairs up to 
4 mm long; margins scaberulous. Panicles 15-30 cm long, 2-6 cm wide, 
mostly open, contracted when immature, pyramidal to ovate; main axis 

Weakley and Peterson, Sporobolus floridanus complex 259 

scaberulous; pulvini in axils of primary branches hairy or glabrous; primary 
branches 2—8 cm long, ascending ro spreading 0—50" from culm axis, not 
floriferous on lower 1/3; secondary branches ascending to spreading; pedicels 
2-22 mm long, usually longer than the spikelet, spreading, scaberulous. 
Spikelets 3-5— 6.5 mm long, purplish-brown. Glumes 2.4-6.5 mm long, 
linear-lanceolate, membranous, 1— veined, subequal to unequal; ratio of lower/ 
upper glume length (0.58-)0.60-0.80(-0.83); lower 2.4-4.5 mm long, apex 
acuminate; upper (3.5-)4-6(-6.5) mm long, apex acuminate to acute. Lemmas 
3.4—4.3 mm long, ovate to lanceolate, membranous, 1-veined, glabrous; 
apex acute. Paleas 3.4—4.4 mm long, ovate, membranous, glabrous; apex 
obtuse to truncate, often bifid with teeth up to 0.4 mm long. Stamens 3, 
anthers 2.5-3.4 mm long, purplish. Grains 1.8-2.2 mm long, fusiform, 

Common name. — Carolina dropseed. 

Distribution and habitat. — Eastern North Carolina south to northern South 
Carolina; and disjunct in eastern Georgia. Wet to moist pine woodlands, in 
soils seasonally to semi-permanently saturated, usually under Pinus palustris 
and Pinus serotina, also sometimes associated with Taxodium ascendens and 
(in Georgia) Pinus elliottii var. dliottii. Sporobolus pinetorum characteristically 
occurs in the following National Vegetation Classification plant associa- 
tions: Pinus palustris-iPinus dliottii var. elliottiij/Sporobolus pinetorum-Aster 
reticulatus—Sporoholus curtissii Woodland; Pinus palustns— Pinus elliottii van elliottii/ 
Styrax americanus var. pulverulentuslSporobolus floridanus Woodland; Pinuspalustris— 
Pinus se)vtina/Sporobolus pinetorum— Aristida stricta-Eryngium integrifolium Woodland; 
Pinus palustr is— Pinus serotinaiSporobolus pinetorum— Ctenium aromaticum—Eriocaulon 
decangulare var. decangulare Woodland; Pinus elliottii var. elliottii— Taxodium 
ascendens /Hypericum brachyphyllum / Sporobolus pinetorum— Dichanthelium scabriusculum 
Woodland (Weakley et al. 1998). July to November (or less typically at 
other seasons if stimulated by fire). 

Comments. — This species has been overlooked until now becatise its over- 
all morphology places it centrally in the complex, allowing it to be vari- 
ously confounded with S. floridanus. S. curtissii, and .V. teretifolius. Despite 
its "intermediate gestalt," it has a unique combination of characters that 
warrant taxonomic recognition. Moreover, it occurs in mixed populations 
with S. floridanus (in eastern Georgia), 5'. curtissii (in eastern Georgia), and 
S. teretifolius (in southeastern North Carolina), and in these situations the 
taxa are easily distinguishable and show no signs of intermediacy or hy- 
bridization. Sporobolus pinetorum is locally abundant in seasonally saturated 
pinelands within its range; because it does not typically flower except fol- 
lowing fire, casual field observers have often overlooked its presence and 
misidentified it as Aristida stricta. 

260 SiDA 18(1) 

Specimens examined. GEORGIA. Chatham Co.: pineland, Ogeechee Canal, 4 Aug 1 939, 
Eylei 6447 (US). Liberty Co.: in ecotone oi: Piii/ts pcih/strislAristichi htyrichicoiii flatwood.s 
and Taxiuiiiim cticcnckns tlrain, 1995, h\ch\'ilhin 1967 (NCU); codominanr wirli SlMmiholiis 
ciirtisiii in mesic longleat pine/saw palmetto tlatwoods on Fort Stewart Military Reserva- 
tion sec. A-5, 4 Sep 1995, McMillan. Kjelhiuirk, & ThoiiipMn 1 171 (USCH); Long Co.: in- 
frec]Lient to rare on mesic longleaf pine savanna and pond cypress depression ecotone on 
Leesfield series soil transition, I'ort Stewart Military Reservation on RSPAC in sect. D-12, 
4 Sep 1995, /VlcAI///rf«, Kjel/mark, & Thow/iwi/ 1 1 70 (USCH). NORTH CAROLINA. Bladen 
Co.: dry savanna 10 mi N of White Lake, 8 Oct 1 944, Blomq/tist / j6/ 7 (DUKE); flat pine 
woods, 6.4 mi ESE of Ammon on road to Garland, 19 Oct 1957, Ahhs 37439 (NCIU). 
Brunswick Co.: low burned savanna on W side ol road to Ec. Caswell, near junction ol 
Sawdust Trail, 1938, Bloiiiquist 1044H (CtH, US); burnt longleaf pine-live oak area near 
inland waterway bridge. Long Beach road, 29 Oct 1950, Boyce & Goilfrey 1602 (NCSC); 
Southport-Supply, savanna, 23 Aug 1930, Bloniqiiht 97 (DUKE, PH); Southport-Supply, 
savanna, 31 Aug 1931, Blomquist 667 2 (YyDKXL, US); Pireway, savanna, 4 Aug \'433,Scbiilleyt 
s.n. (US); shrubby area in pine barrens, Boiling Springs Lakes subdivision, about 2 mi N ot 
jet. of NC 87 & 133, off rte. 133 (N of Southport), 24 Oct 1962, Temll & Smith 3697 
(NCU); plowed fire lane through a bog, Orton, 28 Sep 1941, Godfrey 10053 (NCU). Co- 
lumbus Co.: cut over pine savanna to E ol Co. road, 2.6 mi straight NW of Nakissa [sic; 
probably Nakina], 17 Oct 1958, Bell 13739 (NCU); Schulkens Savanna, 1.7 mi W of Old 
Dock, 17 Jul 1991, Weakley, Schafale. & LeBloml s.n. (NCU, US). Cumberland Co.: Fort 
Bragg Army Base, Macridge Danger Zone, ecotonal seepages and moist swales in upland 
Pinus palustris savannas, 31 Aug 1993, Weakley s.n. (US, NCTJ). Duplin Co.: transition 
near KenansviUe, 14 May 1925, n. c. (NCSC); burned savanna between Clinton and Seven 
Springs, 7 Jul 1946, Blomquist 13936 (DUKE); field woodland border, 0.3 mile NW of 
Pin Hook, 2 Aug 1957 , Ahles & Leisner 33 173 (NCU). Greene Co.: savanna 1.1 mi SE of 
Jason, \95H, Radford 40402 (FLAS). HarnettCo.: l6Nov I992,.S'«me 7/ /6 (NCU). Hoke 
Co.: 7 Oct [991, Sorrie 5945 (NCU); 9 Oct 1991, Some 5959 (NCU); 25 Nov 1 991, Sorrie 
6101 (NCU); 28 Sep 1992, Sorrk 7007 (NCU). Jones Co.: savanna, 1 .2 mi SSW of Pleas- 
ant Hill, 9 Sep 1958, Radford 39935 (NCU); savanna, 3 mi SE of Pink Hill, 9 Sep 1958, 
Radford 39838 (NCU); savanna, 4.3 mi WNW of Hargetts Store, 1 8 Jul 1958, Radford 
36898, (GA, GH, NCU). Lenoir Co.: pine forest, near NC 1 1 , 1 mile N of Pink Hill, 24 
Oct 1957, Radford 31655 (NCU). New Hanover Co.: Wilmington, 1885, McCarthy s.n. 
(US). Onslow Co.: Folkstone, 3 Aug 1939, S/lvens 4856 (US); savanna, 9.6 mi N of HoUyridge, 
27 Jul 1957, Ahles & Leisner 32690 (NCU); Camp Lejeune, 23 Sep 1990, Sorr/e. Weakley. 
LeBlond 5282 (GH); , 5 Sep 199 1 , Churchill 91-170 (BRIT/VDB); cut-over savanna about 
10 mi NW of Holly Ridge toward Maple Hill, common, 1 6 Aug 1 967, Wilh/ir9432 (DUKE); 
WsideofUS 1 7, 0,6 mi S of fire tower, Jun 1991, Weakley & Peel s.n. (\]S). Vender Co.: 30 
Aug. 1991 , Some & LeBlond 5889 (GH, NCU). Richmond Co.: seep in powerline west of 
NC 177, 1 2 Sep 1993, Some 7679 (NCU); Sandhills Game Land, 28 Oct \997 , Sorrie 9621 
(GA, GH, NCU, US). Robeson Co.: woodland border, 2 mi N of Allenton, 2 1 Jun 1957, 
Ahles & Haeslooji 29011 (NCLJ). Sampson Co.: pme savanna, 1.3 mi SE of junction US 
42 1 and 70 1 on US 42 1 (SSE of Clinton), 8 Aug 1951 , Ahles &Lei.uier 33702 (NCU). Scotland 
Co.: 8.3 mi SSE of Eloffman, 1 1 Oct 1959, McNeely 916 (NCU); 4 Sep 1994, Sorrie 8187 
(NCU). [no Co.]: in Oriente Carolina Septentrionalis, locis paludosis [in eastern North 
Carolina, boggy places], Aug \^^5, McCarthy s.n. (NCU, PH, US). SOUTH CAROLINA. 
Berkeley Co.: Jun 1 998, Peet & McMillan 684 1 (NCU). Chesterfield Co.: cleared, burned- 
over shrub bog m the Sandhills 1 mile W of McBee, 7 Sep 1939, Godfrey 8052 (DUKE, 
GH, NCSC, PH, US); savanna, Montrose near US 52, 29 Sep 1956, Radford 18(^(-y7 (NCU); 

Whakley and Peterson, Sporobolus floridanus complex 261 

sandhill, 1 .3 mi SW of" Patrick near US 1 , 29 Sep 1 956, Radford 18761 (NCU); Sandhills 
National Wildlife Refuge, 16 Apr 1992, Some et al. 6246 (USCH); Sandhills National 
Wildlife Refuge, 1995, Pittifian s.u. (US); Hudsoma Flat, 6 Oct 1993, ^rtrr/e 7746 (NCU). 

4. Sporobolus silveanusSwallen,J. Wash. Acad. Sci. 31:350. 1941. (Figs. 
1, g; 3). Type: U.S.A. TEXAS. Orange Co.: 10 mi NE of Orange on U.S. 90, 30 

Sep 1 940, Silvern 644 J (holoiype: US-18 1 7963!; isotwes: US-1867557!, US-2209343!). 

Densely caespitose perennials. Culms 70-1 20 cm tall, erect, nodes sometimes 
visible, mostly basal; base diameter 1 .5^.5 mm, rounded or flattened; internodes 
glabrous. Sheaths mostly glabrous to appressed hairy, hairs up to 4 mmi long; 
base shiny and endurated; margins hyaline; summit with a tuft of hairs, 
hairs up to 4 mm long. Ligules 0.2-0.8 mm long, a line of hairs. Blades 
15—52 cm long, 1—2.5 mm wide, flat to folded or involute, bluish-green, 
glabrous above and below; margins scaberulous. Panicles 21—50 cm long, 
5-12(— 15) cm wide, open and few flowered; pyramidal to ovate; main axis 
scabrous; pulvini in the axils of primary branches glabrous; primary branches 
6-20 cm long, ascending and loosely spreading 20-50? from culm axis, 
not floriferous on lower 1/4—1/2; secondary branches appressed to loosely 
spreading; pedicels 3-8(-l4) mm long, longer or shorter than spikelet, mostly 
appressed, scabrous. Spikelets 4.5-7(-7.2) mm long, purplish. Glumes 3- 
7 mm long, linear-lanceolate to lanceolate, membranous, 1 -veined, subequal 
to unec]ual, ratio of lower/upper glume length 0.6-0.9; lower 3—4.6 mm 
long, apex acuminate; upper 4-7.2 mm long, often appearing 3-veined with 
lateral folds that resemble veins; apex acuminate. Lemmas 4.4-6.5 mm long, 
lanceolate, membranous, 1 -veined, glabrous; margins often hyaline; apex 
acuminate to acute. Paleas 4.5-6.7 mm long, lanceolate, membranous, glabrous; 
apex obtuse to truncate, minutely erose. Stamens 3, anthers 3-5-5 mm long, 
purplish. Grains 1 .8—2.5 mm long, obovoid, laterally compressed, light brownish. 

Common name. — Silveus dropseed. 

Distribution and habitat. — Western Louisiana west to eastern Texas and 
north to southeastern Oklahoma. Wet to mesic pine woodlands under P/>z//j 
palmtris, also in adjoining glade and barren openings, and in blackland prairies, 
5-200 m. Sporobolus silveanus characteristically occurs in the following Na- 
tional Vegetation Classification plant associations: Finns palmtrislSporobolus 
silveanus-IWnhlenbergia capillaris -Liatris pycnostachya var. lasiophylla Wood- 
land; Sporobolus silvmniis-Carex meadit Herbaceous Vegetation; Sporobolus silveanus- 
Tridens strictus Herbaceous Vegetation (Weakley et al. 1998). Flowering 
July to November (or less typically at other seasons if stimulated by fire). 

Comments. — There has been some confusion between this species and S. 
heterolepis. Allen (1992) and Thomas and Allen (1993) reported S. silveanus 
from Calcasieu Parish, Louisiana, and S. heterolepis from Allen Parish and/or 
Calcasieu Parish, but all specimens are actually S. silveanus. Similarly, both 

262 SiDA 18(1) 

S. silvea////s and S. heterolepis have been historically reported from eastern Texas 
(Correll and Johnston 1970; Gould 1975; Johnston 1990; Hatch etal. 1990). 
Brown (1993) determined all material Irom Texas and Louisiana to be S. 
silveanus, and reported S. silveaii/is as a state record for Oklahoma, a conclu- 
sion followed by Taylor and Taylor ( 1 994) and Jones et al . ( 1 997). Sporoboli/s 
silveanus has a bluish color in the field, and closely resembles MNhleiiberg/a 
expansa (Poir.) Trin., with which it often grows. 

Representative specimens. LOUISIANA. Allen Parish: longleaf pine woods, W of Kinder, 
20 Oct 1940, Bnnvn etal. 37/7 (LSLI). Calcasieu Parish: 5 mi S of Starks, 1949, Su./lleu 
105 1 1 (GH). OKLAHOMA. Bryan Co.: pnurie near Durant, 2 1 Oct 1953,.Mftr j.;/. (OKLA). 
TEXAS. Angehna Co.: longleaf pine uplands ol the proposed Graham Creek Wiklerness, 
8.8 mi S of Zavalla on US 69 and E on FR 314, 22 Sep 1 979, N/xon & Ward 9639 (ASTC). 
Brazos Co.: along highway 6, 1 2 mi S of College Station, 8 Oct 1969, Leonard 250 (TAES); 
Galveston Co.: on l4th street one block W of Ave. I (FM 517), 19 Sep 1974, Waller & 
Ba/aiil M2H (SBSC, TAFS, TliX). Hardin Co.: pine forest border, 6 mi SW of Kountze, 
15 Oct 1964, Gould 11028 (BRIT/SMU, TAES, TEX, US). Harris Co.: Red Bluff Road, 
1.5 mi W of SH 146, NW of Seabrook, 18 Sep 1974, Waller & Ba/fwl 3 1 28 (GH, TAES, 
TEX, US). Jasper Co.: SE of Zavalla on US 6i, 1 .6 mi SE of the Plum Ridge Road, 29 
Aug 1978, Alar/e//a & N/xon 486 (ASTC, TEX). Lamar Co.: "Tridens Prairie" 7 mi W of 
Paris at intersection of highway 82 and FR 32, Coll///s .v.//., Nov 197 1 (LL, TAES, TEX). 
Newton Co.: 16 mi N of Newton, 1 1 Oct 1934, Parks & Cory 10832 (TAES). Orange 
Co.: open woods about 10 mi NE ol Orange, 30 Sep I940, S/lveus 6441 (US). Rains Co.: 
in line sandy clay between RR and highway, 3.5 mi NW ol Point, I 2 Sep 1948, Shiiniers 
7(J2.)y (BRIT/SMU). Tyler Co.: longleaf pine grassland, 6.5 mi E of Chester on route 1745 
then left 3 mi to xenc Oligocenc outcrop, 19 Oct 1967, Correll 35 172 (LL). Van Zandt 
Co.: sands. Wills Point, I 5 Oct 1903, Reverchoii 3484 (US). 

5. Sporobolus teretifolius R.M. Harper, Bull. Torrey Bot. Club 3.3:229. 
1906. (Figs. 1, f; 2). Tyim;: U.S.A. GEORGIA: Colquitt Co.: S of Moultrie, 20 
Sep 1902, Harper 1642 (molotvpi;: NY; lsotvpe: US-431954!). 

Caespitose perennials. Culms (20-)35-cSO(-l()0) cm tall, erect, wiry, nodes 
all basal; base diameter 1—2 mm, flattened; internodes glabrous. Sheaths 
glabrous to appressed hairy, hairs up to 4 mm long; base shiny and endurated; 
margms hyaline; summit with a tuft of contorted hairs, hairs up to 4 mm 
long. Ligules 0.2-0.4 mm long, a line of hairs. Blades (I()-)25-54 cm long, 
0.5— 1 .2 mm wide, tightly involute or terete, green to yellowish-green, senescing 
or turning tan in late autumn, glabrous above and below; base often sparsely 
hairy, hairs up to 3 mm long. Panicles 10-26 cm long, 1—9 cm wide, mostly 
open to somewhat contracted when immature, narrowly pyramidal to ovate; 
main axis scabrous; jTulvini in axils of primary branches often hairy; pri- 
mary branches 1-8 cm long, ascending to spreading 0-40" from culm axis, 
not floriferous on lower 1/3; pedicels 3-18 mm long, longer than spikelet, 
usually spreading, with scattered ascending hairs. Spikelets 4—5.6 mm long, 
purplish-brown. Glumes 2—5.6 mm long, linear-lanceolate, membranous, 
1 -veined, unecjual, ratio of lower/upper glume length (0.53-)0.55-0.70(- 

WiiAKLEY AND Petfrson, Spofobolus floridanus complex 263 

0.11); lower 2—3.8 mm long, apex acuminate; upper 4—5.6 mm long, apex 
acuminate. Lemmas 5 A— 4.4 mm long, ovate, membranous, 1 -veined, gla- 
brous; apex acute. Paleas 3-3-4.4 mm long, ovate, membranous, glabrous; 
apex acute. Stamens 3, anthers 1 .5-2.6 mm long, purplish. Grains not seen. 

Common name. — Wireleaf dropseed. 

Distribution and habitat. — Southeastern North Carolina south to south- 
ern Georgia, west to extreme southeastern Alabama. Wet to moist pine 
woodlands, \xx\Att'Pinus serotina. Pinus elliottii var. elliottii, and Pinus palustris, 
in soils; 10—150 m. Sporobolus teretifolius characteristically occurs in the fol- 
lowing National Vegetation Classification plant associations: Pinus palustris— 
Pinus serotina/ Magnolia virginianalSporobolus teretifolius— Carex striata 
Woodland (Weakley et al. 1998) and others not yet described. Flowering 
July to November (or less typically at other seasons if stimulated by fire). 

Comments. — The terete leaves (well figured by Harper 1906) are distinc- 
tive. Dry blades (either on dried specimens or in nature under dry field conditions) 
of the other eastern species with narrow blades (5'. curtissii. S. pinetorum) can 
superficially resemble those of 5'. teretifolius, and have caused confusion. Sporobolus 
curtissii, however, has much shorter blades and generally occurs in drier habitats, 
and the blades of S. pinetorum have distinctly rough margins (best felt by 
running ones finger along the blade towards its base, or seen at 10" or greater 

Representative specimens. ALABAMA. Houston Co.: James Hughes pitcher plant bog, 
1.5-2.0 mi NE of Cottonwood, burned annually, 4 Aug 1996, MacDomild 9833 (IBE). 
GEORGIA. Berrien Co.: very broad powerline on N side of route 76, about 1.7 mi SW of 
Nasliville, 27 Oct 1 994, Sorrks.n. (NCU). Bulloch Co.: sloping moist pine barrens, about 
7 mi NW of Statesboro, 12 Sep 1954, Harper 4309 (GA). Candler Co.: about 3.25 mi W 
of Matter on Stillmore Road, ecotone on E side of a N-S swampy tributary of Sams Creek, 
S side of road, 22 Apr 1994, Some & Stoireli s.n. (NCU). Coffee Co.: rather dry pine-bar- 
rens near Douglas, upper Eocene overlaid by Lafayette and Columbia, 22 Sep 1900, Harper 
677 (NY). Colquitt Co.: moist pine barrens near Moultrie, 20 Sep 1902, Harper 1642 (G A, 
NY, EJS. Cook Co.: low seepage area (pitcher plant bog) alongside GA 76, 2.7 mi NE of 
Barney, 2 1 Sep 1 965 , fainioth & O'Neal 2994 (GA). Dodge Co.: reported without precise 
location by Harper (1906). Dooly Co.: reported without precise location by Harper (1906). 
Emanuel Co.: moist mown margin of GA 57 just NW of milepost 5, S side of road, at 
edge of shrubby ecotone of the W side of Flat Creek, NW of Swainsboro, 6 May 1994, 
Sorrie s.n. (NCU). Screven Co.: E side of US 301 at South Fork Ogeechee Creek (milepost 
7), S of Sylvania, 25 Oct 1994, Sorrie s.n. (NCU). Thomas Co.: Greenwood Plantation, 
Thomasvillc, damp peaty soil bordering a Magnolia virginiana—Nyssa hiflora drainage, area 
burned June 1985, 23 Oct 1985, Gholson, Godfrey, Komarek. & Baker 11522 (GA). Tift 
Co.: Vicinity of Irby P.O. (Cycloneta Station), 28 Jul 1890, Tracy s.n. (NY) {note that present 
Tift Co. was part of Irwin Co. at the time of Tracy's collection}. Toombs Co.: among a 
colony of pitciier plants, 3 mi N of Lyons, 28 Jul 1961, Banks s.n. (GA). Turner Co.: seep- 
age slope flanking route 32, W side of Little Sand Creek, 25 Aug 1994, Sorrie s.n. (NCU). 
Wheeler Co.: moist pine barrens, 2 mi W of Alamo, Plummer sandy loam, Pullen & Pliitmner 
881 (GA). NORTH CAROLINA. Brunswick Co.: Camp Branch Savanna remnant, just 
N of SR 1335, ca. 1.2 mi E of its junction with SR 1334, back edge of savanna, near fire 

264 Si DA 18(1) 

(.lirch at c'cocone CO j-iocosiny swamj-i, K\)ul 1991, Wtakk']& Schiiliile s.ii. (US). Columbus 
Co.: Old Dock Savanna, south ol SR 1928, ca. 0.9 mile W of Old Dock, then south on 
logging road ca. O.t mi, 15 Jul 1991, Wt'akky & Schajalc s.n. (US). SOUTH CAROLINA. 
Georgetown Co.: pine savanna in Bates Hill Plantation, ca. 4.0 km SE of the US 701/SC 
261 intersection, 5 Sep 1989, 'I'iii^^ii^iirt s.ii. (USCH). Horr)' Co.: along powerline, situated 
between Mose Swamp anti Grass Bay, take SC 1 09 ca. 2 mi NW oi Baker Crossroads, then 
W and intersect with Sancee Cooper Power right-obway, 22 Sep 1992, Pittman & Jordan 
s.n. (USCH). Kershaw Co.: mesic slope, SC 28-102, 2.8 mi N of SC 12, 29 Sep 1958, 


Stapf ( 1 89<S), Bor (I960), Clayton er al. (1974), and Baaijens and Veldkamp 
(1991) have suggested infrageneric classifications in Sporoholiis based pri- 
marily or strictly on Old World species, and a more worldwide classifica- 
tion is still lacking. Based on possession of an open or contracted panicle, 
caespitose perennial lifeform, non-whorled branch insertion, intravaginal 
new shoot initiation, and first gltime shorter or as long as the second Pilger 
(1956) erected group four in subgenus Sjiorohoh/.s. He further divided group 
four into two smaller groujis a cs: B. Sporoholiis beterolepis, S. floruhnuis. S. 
teretifoluis. and S. lasiophyll/is Pilg. are members of Pilger's "B" group. We 
agree with Pilger's assessment that.V, heterolepis. S. floridanus, and S. teretifoPiiis 
appear to be closely related. However, S. Lisiophylh/.s lias a few unique char- 
acteristics, such as its strictly basal leaves with sheath bases that are densely 
lanate and thickened, and its plumbeous to dark-brownish spikelets. 

Small (193.3) divided Sporoholiis of the southeastern United States into 
seven groups, which are not considered validly published because their rank 
wiis not indicated. He placed S. jiinceiis [iis ".V. gracilis (Trin.) Merr. "], S.florickiniis, 
S. ciirtissii, and S. teretifoluis in '^Grcuiles. " distinguished by perennial, bunchgrass 
habit, large (more than 3 mm long), purplish to brown spikelets, and glumes 
unequal, the second as long as the floret. Sporoholiis heterolepis was not known 
by Small to occur in his "flora area," so it is unknown whether he would 
have placed this with ''Graciles" or not. 

Wc informally recognize the S. floridaniis complex, consisting of five species 
(.V. ciirtissii. S. floridaniis, S. pinetoriim, S. silveaniis, and S. teretifoliiis) with 
veiy similar morphological features and generally similar ecological requirements 
(primarily restricted to pine savannas on the Coastal Plain). All five species 
exhibit the following characteristics: caespitose perennials; basal sheaths shiny 
and endurated (apparently as "fire-proofing"); panicles open at maturity, 
with ascending panicle branches, pyramidal to ovate, primary branches not 
floriferoLis on lower 1/3; spikelets purplish-brown to purplish; lower glume 
linear-lanceolate to lanceolate, apex acuminate; grain fusiform to obovoid. 
Additionally, all five species occupy fire- main rained pinelands of the southeastern 
Coastal Plain, and produce culms only following removal of foliage by fire 

Weakley and PetersoiN, Sporobolus floridanus complex 265 

(or rarely, by mechanical disturbance) [Weakley 1998}. 

The S. floridan//s complex appears to have its closest affinities with two 
pairs of species: 1. S. heterolepis and S. interriiptiis Vasey, and 2. S. jiinceiis and 
S. purpurascens (Swartz) Hamilton. Sporohh/s heterolepis and S. interrupt/is ap- 
pear to be sibling species, S. heterolepis being widely distributed in central 
North America, and extending as an uncommon disjunct into eastern North 
America, and S. interrupt //s being endemic in northern Arizona. They share 
several characters which distinguish them from the S. floridanus complex: 
spikelets plumbeous (vs. purplish fading to tan), grains globose (vs. elon- 
gate and laterally flattened), scaberulous lower glumes (vs. glabrotis), and 
fibrous sheath bases (vs. shiny and indurated). Sporobolus heterolepis appears 
to be particularly closely related to S. silveanus, the most western of the S. 
floridanus complex, and the one most likely to have been geographically 
and ecologically in contact with S. heterolepis in recent times. Ecologically, 
S. heterolepis is a species of glades, prairies, and barrens, usually overmafic, 
ultramafic or calcareous rock outcrops in the interior, "hard rock" physi- 
ographic provinces of central North America (extending as a rare disjunct 
east to and occasionally beyond the Blue Ridge Mountams), as well as on 
loess and glacial tills. 

Sporobolus junceus and S. purpurascens also appear to be siblings, S. junceus 
being a species primarily of the southeastern United States Coastal Plain 
(from southeastern Virginia west to eastern Texas), but extending inland to 
adjacent provinces, while S. purpurascens is distributed in southern Texas, 
Mexico, the West Indies, and into tropical America. They share several characters 
which distinguish them from the S. floridanus complex: panicle branches 
distinctly whorled in well-marked verticils (vs. panicle branches alternate 
or sometimes irregularly paired or approximate) and spikelets smaller (3-0- 
3.8 mm long vs. 3.5-7.2 mm long). 


Sporobolus curtissii, S. floridanus, S. pinetorum, S. silveanus, and S. teretifolius 
are all relatively narrow endemics of various portions of the southeastern 
Coastal Plain (Figs. 2, 3). Each of these five species is the locally dominant 
or codominant grass in fire-maintained pinelands with open canopies oiFinus 
palustris, P. serotina, and/or P. elliottii var, elliottii (Weakley et al. 1998). The 
geographic distributions and many ecological requirements of the four more 
eastern species overlap, but they can be separated based on a hydrologic gradient. 
Understanding the differing but overlapping distributions and ecological 
niches of the species within the group provides an important basis for the 
systematic treatment. 

The five species of the complex share a set of apparent adaptations to the 

266 SiDA 18(1) 

fire-maintained habitats in wliich they ocx-tir. Ah have the basal sheaths thickened 
and cartilaginous, tightly investing the growing tips of the rhizomes, and 
protecting it from damage or destruction by fire. This is a conspicuous fea- 
ture of herbarium specimens (if not removed by overzealous preparation), 
appearing as a stramineous, shining, thickened (almost bulbous) base of the 
plant. The bases of Calamovilfa brevipiln and Calamovilfa cun'mit (Vasey) Scribn. 
are very similar. Species of the S. floridanus complex re-sprout quickly fol- 
lowing fire, with green leaf material protruding beyond the blackened basal 
sheaths within days following fire. Like other southeastern bunchgrasses 
adapted to fire-maintained pinelands, including Calamovilfa hrevipilis. C. 
ii/i'tissii. Aristida stricta. A. beyrichiana. and Ctenii/ii/ aro»/atu'//>!J, all fwe Sporobol/zs 
species generally produce culms only after having their leaves removed by 
fire, although they will sometimes flower in response to mechanical distur- 
bance. For this reason, nearly all herbarium specimens have the upper por- 
tions of the basal sheaths conspicuously blackened. 

General exclusion of fire from much of the pineland habitat of the five 
species of the S. floridanus complex, combined with their flov/ering only in 
response to fire, means that the species are often overlooked or misidentifled, 
and that they are severely under- represented in herbaria despite their local 
abundance. The four narrower-bladed species, S. pmetornn/, S. ciirtissit, S. teretifoli/is. 
and S. silveaN//s, are often actively or passively misidentifled as other sym- 
patric pineland bunchgrasses, ^uc\\ 'j&Anstidastncta, A. beyrichiana, or Muhlerihergta 
expansa. Entire savannas of a hundred hectares or more dominated hyS. pinetorum 
have been assumed to be "longleaf pine/wiregrass," thotigh in reality Aristida 
stricta may be completely absent. For this reason, it is important that field 
biologists become familiar with the vegetative characters which allow rec- 
ognition of these taxa in sterile condition. 

The complex reaches its greatest diversity in the Coastal Plain of C^eor- 
gia, whf^tt S.floridan//s. S. airtissii. S. teretifolius, and S. pinetoriini are all found, 
and co-occtir in various combinations of two and three species, along with 
the related S. juucciis. Sporoboh/s silvcaiuis is the only species of the complex 
that is fully allopatric, and the only one that occurs west of the Mississippi 
River. In southeastern North Carolina, S. pinetor/nii and S. teretifdias occur 
in wet pinelands dominated by mixtures oi Piniis palustris, Finns serotina, 
and Taxodimn ascendens. They generally co-occur with other savanna bunch- 
grasses, notably Cteniumaromaticuni. Calamovilfa brevipilis, M//hlenbergia expansa, 
and Aristida stricta. Sporoboliis teretijolij/s is restricted to the wettest pine sa- 
vannas, usually so wet as to exclude Aristida stricta, and it may be the sole 
dominant, or codominant with S. pinetorum. Ctenii/m aroniaticiim, and Muhlenbergia 
expansa. Slightly less wet savannas have varying mixtures of 5", pinetorum, 
Aristida stricta. Muhlenbergia expansa, and Cteniiim aroma tici/ni. A similar 

Weakley and Peterson, Sporobolus floridanus complex 267 

composition, though often with substantial Calamovilfa hrevipilh as well, is 
seen in sandhill/pocosin ecotones in the inner Coastal Plain. 

A general hydrologic gradient of the five species and the sympatric S. 
junceus would be (from wetter to drier) S. floridanus. S. teretifolius, S. pinetorum. 
S. silveanm, S. curtissti, S. junceus. Sporobolus floridanus is restricted to habi- 
tats which are saturated at least seasonally (and often semi-permanently) 
and may even have shallow ponds for weeks or even months. The other spe- 
cies are generally intolerant of ponds for more than a few days. Sporobolus 
teretifolius occurs in habitats with semi-permanent to seasonal saturation, 
whereas S. pinetorum occurs in habitats with seasonal saturation. Sporobolus 
stlveanus and S. curtissii occur in a range of sites, from seasonally saturated 
to sites which are rarely saturated at the surface for periods of short dura- 
tion only, and may range up to rather well-drained (though not xeric) sites. 
Sporobolus junceus strictly occupies well-drained sites, ranging into truly xeric, 
sandhill situations, where it reaches its greatest abundance. 

Sprobolus floridanus generally occupies the wettest habitats of the com- 
plex. Particularly towards the edges of its distribution, it is generally found 
in very wet situations, often where water stands for periods of time, often 
in seepage bogs or swales, and generally associated with Pinus elliottii var. 
elliottii and Taxodium ascendens. Near the center of its distribution, especially 
in the eastern portion of the Florida panhandle, S. floridanus ranges into 
less wet habitats, and occurs in "mcsic flatwoods." Where its distribution 
overlaps with 5", pinetorum. S. curtissii, and S. teretifolius, it can occur (with 
clumps side by side), but it clearly ranges ecologically into wetter sites and 
avoids drier sites. 

The four eastern species in the S. floridanus complex commonly co-occur 
in pineland landscapes, and sometimes occur in mixed populations, with 
individuals of various species intermixed. In these circumstances, the vari- 
ous species are always readily distinguishable by morphological characters. 
No intermediates or likely hybrids have been seen. None of these species 
has been investigated cytologically. 


We wish to thank Michael P. Schafale, Richard LeBlond, Bruce A. Sorrie, 
Brian van Eerden, Harry LeGrand, Jr., Patrick MciVIillan, and Robert K. 
Peet for discussions regarding this group and for providing additional dis- 
tribution information; Susan C. Escher for preparing the illustration; Alice 
Tangerini for preparing the distribution maps; Victoria E. Batista for pre- 
paring the Spanish abstract; Dan H. Nicolson for help preparing the Latin 
diagnosis; Dieter C. Wasshausen and Gene Rosenberg for help translating 
German; the curators of the listed herbaria, and especially Jim Massey at 

268 Si DA 1,S(1) 

NCU for providing facilities to the first author; and David W. Hall, Stephan 
L. Hatch, and Robert K. Peet for critically reviewing the manuscript. 

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Department oj Biology 

Washington University 

Campus Box 1 137 

St. Lonis. MO 63130. U.S.A. 

P.O. Box 299 

Missouri Botanical Garden 

St. Louis, MO 63166. U.S.A. 


Caryopsis morphology of the grass genus Leptochloa was studied regarding the necessity 
of some authors for segregating Dtplachne. The data do not support the separation of Dipliulme 
from Leptochloa based on a putative bimodal distribution of dorsal and lateral cross-sec- 
tional compression. The presence or absence of a prominent sulcus and the relative achia- 
tion of the pericarp are the only attributes sufficiently distinct to warrant use as phyloge- 
netic markers. However, variations in surface texture and color can be useful regionally as 
diagnostic characters at the species level. 

Key worixs: caryopsis, Ltptochloa, Diplachne, morphology, systematics, Poaceae, Chloridoideae. 


Se estudio la morfologia del cariopside del genero Leptocloloa dada la supuesta necesidad 
de algunos autores de segregar Diplachne. Los datos no apoyan la separacion de Diplachne y 
Leptochloa basada en una supuesta distribucion bimodal de la compresion dorsal y en seccion 
transversal lateral. La jiresencia o ausencia de un surco prominente y la adnacicMi relativa 
del j->ericarpo son los unicos atributos suficientemente diferentes para justificar sti uso como 
marcadores filogeneticos. Sin embargo, las variaciones en textura y color de la superticie 
pLieden ser utiles regionalmente como caracteres cliagnosticos a nivel especitico. 


The genus Leptochloa P. Beauv. s.l. (including Diplachne P. Beauv.) has been 
the subject of numerous regional systematic studies due to its wide geo- 
graphic distribution and the relative abundance of herbarium specimens 
(Hitchcock 1903; Parodi 1927; McNeill 1979; Lazarides 1980; Phillips 1982; 
Nowack 1994; Nicora 1995). These authors (and others) have disagreed as to 
whether Diplachne should be segregated from Leptochloa, thereby mirroring 

'Ctirrent Address: Department of Biological Sciences, University of Northern Colorado, 
Greeley, CO 80639, U.S.A. Email; nsnowCf; 

SiDA 18(1): 271-282. 1998 

272 Sum 18(1) 

the ditterini,^ opinions of two worldwide generic summaries of grasses (Clayton 
& Renvoize 1986; Watson & Dallwitz 1992). A frequently cited source of 
evidence to support the segregation of D'tplachm has been differences in caryopsis 

Parodi (1 927) apparently was the first to examine cross-sectional shapes 
critically. He partitioned four neotropical species into Diplachne or Leptochloa 
based on relative compression (dorsal or lateral) of the caryopsis, and the 
presence or absence of a hilar groove. He suggested that Leptochloa chloridiformh 
was aberrant in Leptochloa because of its lack of a hilar groove (Parodi, I.e.). 

Vails (1978) studied the systematic affinities oi Leptochloa dubia in rela- 
tion to the generic boundaries of Leptochloa. He illustrated cross-sectional 
profiles and profiles from the embryonic and hilar sides for seven species. 
The figures revealed a gradation in cross-sectional profile from nearly round 
in Diplachne caitclata to somewhat triangular in Leptochloa virgata and L. scabra, 
to relatively Battened in L. fascicz/laris (Vails, I.e.: 103). He also concluded 
that a hilar depression was a tenuous systematic feature. Despite somewhat 
limited sampling, his results suggested strongly that caryopsis features intergrade 
too thoroughly to split Leptochloa unambiguously into two genera, although 
he acknowledged that "some grouping of species can be achieved on the 
basis of caryopsis type" (Vails, I.e.: 105). Unfortunately, his results were 
never formally published and have not been cited by subsequent authors. 

McNeill (1979: 401) and Nicora (1995: 233) repeated almost verbatim 
the observations of Parodi (1927) without adducing additional data or cit- 
ing the work of Vails ( 1 978). Lazarides ( 1 980) observed that Australian species 
generally could be segregated into Leptochloa or Diplachne on the basis of 
caryopsis siiape, with the exception oi^ Leptochloa digitata, with its flattened 
shape. Phillips (1982: 144) agreed with Parodi (1927) regarding the util- 
ity of caryopsis shape for splitting the genera, but noted exceptions in Diplachne 
caudata, Leptochloa obtusiflora, and L. longa. Nowack (1994) provided a cur- 
sory review of caryopsis shapes for IMalesian taxa and concluded that the 
differences set forth by Parodi (1927) were insufficient to permit recogni- 
tion of segregate genera. 

Based on a recent monographic treatment (Snow 1997), Leptochloa (in- 
cluding Diplachne) represents forty taxa, with one species, L. monticola Chase, 
being of dubious inclusion (Vails 1978; Snow 1996). Prior to this study, 
relatively few taxa had been examined critically for variation in features of 
the caryopsis, and much systematic weight had been placed on the meagre 
observations that existed. In light of lingering debates about generic boundaries 
between Leptochloa and Diplachne (Jacobs 1987) and the emphasis previous 
authors placed on the utility of the caryopsis to segregate these genera, a 
survey of all currently recognized taxa of Leptochloa was undertaken to evaluate 
whether features of the caryopsis could be useful as phylogenetic markers. 

Snow, Caryopsis morphology of Leptochloa 273 


Caryopses of all currently recognized taxa in Leptochloa (Snow 1997) were 
removed directly from herbarium specimens (Appendix 1), placed under a 
Nikon SJMZ-U dissecting microscope with camera lucida attachment, and 
the profiles were traced by hand. Cypholepis yemenicus was included because 
it resembles L. eleusine and L. obtusiflora in several respects (Snow 1996), 
and was used as an outgroup in preliminary cladistic studies o^ Leptochloa 
(Snow 1997; see also van den Borre & Watson 1997). In most cases a mini- 
mum of three specimens were examined for variation (Appendix 1). The 
following features were observed: 1) caryopsis shape when viewed from the 
hilar side ("hilar profile"); 2) caryopsis shape when viewed from a cross-sec- 
tion taken at midpoint with the hilar side oriented above ("cross sectional 
profile"); 3) the presence or absence of a sulcus or other depression on the 
hilar side when viewed in cross-section; 4) ornamentation on the outer coat 
(perisperm); 5) relative adnation of the perisperm to the endosperm, and 6) 
color of the grain. To standardize the sampled developmental stage, cary- 
opses were selected from spikelets in which florets were beginning to disar- 
ticulate, a condition that assures their maturity. In virtually all cases the 
caryopses from the lowermost floret in the spikelets were selected. 

For the sake of precision, descriptive terminology of shapes follows that 
of the Systematics Association (1962), whereas that of surface ornamenta- 
tion follows Murley (195 1 ). Given that shape is a continuously varying character, 
these typologies might not account for the observed and often subtle varia- 
tions in shape. For example, a caryopsis might have an intermediate ellip- 
tic shape of 2.5:1, which is absent from the diagram. Nonetheless, after 
initial analyses, the diagram shapes appeared adequate to standardize and 
summarize the majority of both hilar profile and cross-sectional shapes. With 
respect to cross-sectional shapes, I accounted for the absence of sharp edges 
by prefixing the terms "obtriangular" (3:2), "shallowly obtriangular" (3:2), 
and "shallowly obdeltate" with the word "rounded," which more accurately 
depicts their shapes. To account for concave inflections of the hilar surface 
(always oriented above, Fig. 2) I used the terms "sulcus" and "depression," 
depending on the degree of concavity. As used here, a sulcus is a vertical or 
nearly vertically walled groove; a depression refers to any gradual concav- 
ity, and will be further modified by the terms shallow, moderate, deep, narrow, 
and broad. These subjective terms were deemed necessary to describe the 
observed variation. 


The hilar profile for most species was some variation ot obovate or ellip- 
tic (Fig. 1; Table 1). Only four taxa had ovate hilar profiles {L.fusca subsp. 


SlDA 18(1) 

Narrowly elliptic 










2:1 3:2 

Widely elliptic 


Widely obovate Very widely obovate 



Fig. 1. Hilar profiles of caryopses observed tor LepUxhioa. Tlie apex is oriented above. 

Snow, Caryopsis morphology of Leptochloa 


Table 1. Variation in hilar profiles and cross-sectional shapes of the caryopsis in l^eptochloa s. 1. The 
numbers following each taxon reflect observed variation: numbers to the left of the double bar (|j) 
indicate hilar profile shapes (see Fig. I); those to the right of the double bar reflect cross-sectional 
shapes (Fig. 2).. Hence, 2,3,8 || 2,9 would indicate a taxon having narrowly elliptic (.3T), elliptic 
(2:1), and obovate (2:1) hilar profiles, and oblate and depressed obovate (2:.3) cross-sectional shapes. 
Taxa with an asterisk (*) indicate those previoLisly placed in Dtplachm by some authors. 

Leptochloa at]uatica .3,9,10 || 8,9,12 

L. caudata*l,2,8 || 2 

L. chinensis 8,9 || 2,9 

L. chloridiformis 2 || 7 

L. coerulescens 8,9 || 2,8 

L. decipiens subsp. asthenes 3,8 [| 2,8,9 

L. decipiens subsp. decipiens 3 |[ 9 

L. decipiens subsp. peacockii 3,8 || 3,9 

L. digitara 2,3,8 1| 8,12 

L. divaricatissima 3 j| 2,9 

L, dubia* 2,3,11 || 8 

L. eleusine* 3,8,9 || 8,12 

L. fusca subsp. fascicularis* 3,8 j] 4 

L. fusca subsp. fusca* 9 j| 4,8 

L. fusca susbsp. muelleri* 3,9 || 4,8 

L. tusca subsj-). uninervia* 3,7,8 jj 4 

L. gigantea* 3,6,8 || 8 

L. longa 3,8 || 9 

L. ligulata 3,4 jj 2,7 

L. malayana* 3 || 9,11 

L. marquisensis 2,3,6 jj 9,10 

L. monticola* 3,8 || 4,8 

L. nealleyi 4,9 || 2,9 

L. neesii 5,10 j| 1 

L. obtusiflora 3 j| 8 

L. panicea subsp. brachiata 3,4 || 10,13 

L. panicea subps. mucronata 3 |j 1,2,8 

L. panicea subsp. panicea 7 [| 1,2 

L. |ianicoides* 3,4 j| 8 

L. rupestris 3 || 10 

L. scabra* 2,3 jj 9 

L. southwoodii 3,4,8 jj 1,2 

L. squarrosa 2 j| 5,11 

I., srilankensis 3 jj 9,10,11 

L. tectoneticola* 2 j| 8 

L. uniflora 2,6 jj 6,7 

L. virgata 2,3,6 j| 6,7,1 1,12,13 

L. viscida* 3,8 II 4 

L. xerophila 3 jj 2 

Cypholepis yemenicus 4 j| 8,9 

unimrvia^ L. gigantea, L. uniflora, L. virgata). The widest was the very widely 
elliptic shape of a few specimens of L. dubia. The thinnest was the narrowly 
elliptic (3T) shape expressed by some specimens ofL. caudata, L. chloridi- 
formis, L. dubia, L. digitata, L. scabra, L. squarrosa, L. uniflora, and L. virgata 
(the lattermost sensu lato, including L. barbata and L. procera sensu Nicora 
1995). Many species were variable, for example having both elliptic (2:1) 
and obovate (2:1) shapes. Not surprisingly, the greatest variation in hilar 
profile shape occurred in widespread species such as L. dubia and L. virgata. 
The cross-sectional shape was considerably more variable than hilar profile 
shape (Fig. 2). Overall, the observed variation ranged from dorsally com- 
pressed through circular (no compression) to laterally compressed (Fig. 2). 
Most taxa had only slight to moderate degrees of lateral or dorsal compres- 
sion. As with hilar profiles, many taxa showed infraspecific variation in cross- 
sectional shapes (Table 1). Some specimens of L. neesii appeared circular, 
whereas others were oblate (Table 1). The greatest degree of dorsal com- 
pression was expressed by the depressed obovate (1:2) and transversely el- 
liptic (1:2) shapes. With some modifications, these shapes accounted for 
some or all of the variation of many species (Table 1). The highest degrees 
of lateral compression were the obovate (3:2), rounded shallowly obdeltate 

276 SiDA 18(1) 

(5:6), and rounded shallowly obtriangular shapes. Only L. squarrosa was obovate 
(with a moderate hilar depression), whereas some representatives of L. virgata 
were both rotinded shahowly obdeltate (5:6) or rounded shallowly obtriangular 


A distinct sulctis was present only for L. n/pestris and L. uniflora, although 
a number of taxa had depressions of varying extent on the hilar surface (see 

The surface of the pericarp varied from smooth to variously rugose. The 
following were at least occasionally somewhat rugose: L. chloridifornt'n, L. 
ilecipieiis subsp. decipiens, L. divciriccitissiDUi, L. g/gaiilea, L. hmga, L. //udayai/a, 
L. riioiitu'olci, L. riealleyi, L. neesii, L. obtus/flora, L. saibra, L. soi/thwoodii, L. 
uniflora. Whereas a smooth pericarp was consistent for many taxa, those that 
expressed the rugose condition did so irregularly. 

Species with a weakly adnate pericarp (detaching soon after placement 
in water at room temperature) included L. chloridiforniis, L. duhia, L. e/e//sh?e, 
L.fusai subspecies ///icv/, fascicnhiris, un!)iervia, and L. obtusiflora. 

The color of the caryopsis varied from very light brown to dark reddish 
or very dark brown, but most were an intermediate shade. Leptochloa longa, 
L. obt//sJflora, and L. sqiiarrosa were tisually dark brown. Leptochloa monticola, 
a species of dubiotis inclusion in the genus (Clayton & Renvoize 1 986; Vails 
1978; Snow 1996, 1997), was usually a dark reddish brown. 


This simple study of caryopsis morphology has revealed more variation 
within and between taxa o( Leptochloa than previously recognized (Parodi 
1927; McNeill 1979; McVaugh 1983; Nowack 1994; Nicora 1995). With 
two exceptions, features of the caryopsis appear to be of little value in Leptochloa 
as phylogenetic niarkers, although some are of diagnostic value in keys. These 
results contrast with those of a recent study in Triticeae, which suggested 
caryopsis morphology was of systematic value at the tribal level (Terrell & 
Peterson 1993). 

Parodi's ( 1927) study was limited to four species in Leptochloa and one in 
Goutnia Fourn., which represents only ten percent of Leptochloa as currently 
circumscribed (Snow 1997). Another disconcerting aspect was his lack of 
voucher specimens and tmcertain depth of sampling within taxa, although 
this study does not contradict the profiles of the species he illustrated. Overall, 
Parodi's sampling underestimated considerably the variation in cross-sec- 
tional shape in Leptochloa. This study has revealed nearly continuous varia- 
tion in cross-sectional shape, from dorsally compressed and non-compressed 
(circular or oblate) to various degrees of lateral compression. The bimodal 
compression (lateral or dorsal) ofcaryopses in Leptochloa recognized loy Parodi 

Snow, Caryopsis morphology of Leptochloa 





Transversely elliptic 





Obovate Widely obovate Very Widely obovate 



Depressed obovate 


Widely depressed obovate 





"Rounded" "Rounded" shallowly "Rounded" shallowly 
obtriangular obtriangular obdeltate 





Fig. 2. Cross sectional sliapes of caryojises observed tor Leptochloa. The hilar side is oriented 

278 SiDA 18(1) 

(1927) simply does not exist for the genus as a whole. As such, variation in 
caryopsis cross-sectional morphology cannot by itself be invoked as a ge- 
neric-level character to segregate Diplachne. 

The caryopsis profile as viewed from the hilar side ranges continuously 
from narrowly elliptic through ovate to obovate and very widely obovate 
(Fig. 1 ). Whereas the extremes ot variation can be useful as diagnostic fea- 
tures between some taxa, the continuous variation makes the hilar profile 
useless as a phylogenetic marker (Stevens 1991 )• 

Various degrees of concavity occur on the hilar side. Broad, shallow de- 
pressions (not illustrated) characterize certain taxa fairly well, such as Leptochloa 
eleusine, L. ionga, and L. obtustflora. Taxa showing this feature irregularly were 
Leptochloa chinensis, L. di/bia, and the related Cyphokphis yernenicns . Shallow, 
relatively narrow depressions (not illustrated) occur in other taxa, althotigh 
less consistently; these included L. chloridijormis, L.decipiens subsp. decipiem^ 
L. dtgitata, L. sqiMrrosa, and L, virgcita. At best, the degree of concavity is 
useful only as a diagnostic character in regional keys. However, a promi- 
nent sulcus, characterized by its vertical or nearly vertical walls, was a con- 
sistent character for L. rnpestrh and L. uniflora, and is one of only two char- 
acters I consider sufficiently distinct and consistent to be phylogenetically 

The relative adnation of the pericarp is the second character of the cary- 
opsis useful for phylogenetic inference. It is well known that the pericarp is 
only weakly adnata to the endosperm in some species of Leptochloa (Izaguirre 
& Laguardia 1987; Watson & Dallwitz 1992) and some related genera, such 
as Eragrostis Wolf (Lazar ides 1997). In such taxa the pericarp will dissociate 
from the endosperm quickly when placed in water at room temperature. 

iVIost species in Leptochloa have a smooth outer texture. A few can be rug- 
ose, althotigh this feature was unreliable within taxa. For example, a coarse 
but sparsely rugose surface generally, but not always, characterizes Leptochloa 
pantcea stibspecies panicea and mucronata (sensu Snow 1 99Ba, but not sensu 
Nowack 1994), which helps to distinguish these from the widespread L. 
panicea subsp. brachiata (Snow 199Ha; formerly known as L.jilifornih or L. 
mucronata [Snow & Davidse 1993}). 

As the color of the caryopsis often varies with the degree of maturity, 
only mature specimens should be evaluated for this attribute. Except as a 
diagnostic feature in keys, in which a few species are dark brown, color is of 
minimal systematic value in Leptochloa. 

I return now to cross sectional shape, which has been discussed exten- 
sively regarding the separation o'i Diplachne from Leptochloa (Parodi 1927; 
McNeill 1979; Phillips 1982; Nicora 1995). As mentioned above, the per- 
ceived bimodality of lateral and dorsal compression disctissed by Parodi (1927) 

Snow, Caryopsis morphology of Leptochloa 279 

has been invoked to segregate Diplachne from Leptochloa. The results of this 
study firmly reject such a notion, given the nearly continuous variation of 
cross-sectional hilar profiles (Fig. 1). 

It also has been suggested that the presence or absence of a distinct keel 
on the lemma is positively correlated with cross-sectional shape, and is a 
means by which the genera can be separated (Parodi 1927; McNeill 1979; 
Nicora 1995). It is true that some taxa with a dorsally compressed cary- 
opsis have flat lemmas at maturity, as for example L. fusca subsp. muelleri. 
However, others have little or no such positive correlation. Mature fruits of 
L. neesii can be round or nearly so in cross section, yet still be borne within 
a keeled lemma (e.g. , Lan(^ld283 , CANB). This is also true for some specimens 
of L. ligulata, L. nealleyi, L. pankea subsp. panicea, and L. southivoodii. Pre- 
liminary cladistic studies have failed to consistently group together taxa 
having dorsally flattened caryopses (Snow 1997). 

This study upholds and strengthens the unpublished work of Vails (1978), 
who concluded that variation in caryopsis shape was too great to support 
the recognition of Diplachne. In particular, the data herein have revealed 
nearly continuous variation in cross-sectional shape, ranging from dorsal 
compression through circular to various degrees of lateral compression (Table 
1; Fig. 2). The lateral/dorsal compression dichotomy of Parodi (1927) sim- 
ply does not exist for Leptochloa. Moreover, a nearly identical range of cross- 
sectional shapes can exist in closely related genera, as illustrated in a recent 
revision of Australian £r^^^roj//j- (Lazarides 1997: 176). 

The most general observation to emerge from this and other detailed anatomical 
and morphological studies of grasses (Davila & Clark 1990; Ellis & Linder 
1992; Snow 1996; Lazarides 1997) is that broader sampling regimes gen- 
erally reveal additional variation not encountered in narrower surveys. Such 
variation cannot be ignored. Future studies therefore should seek both breadth 
and depth of sampling in order to minimize the chances of incompletely 
characterizing variation. Underestimates of variation leads to errors in the 
diagnosis of taxon boundaries and in the accuracy of inferring phylogenetic 
relationships, the latter being the very basis by which we make and sup- 
port our classifications. 


Generous funding was provided by the National Geographic Society (NGS 
5594_95) and the Missouri Botanical Garden (Andrew W. Mellon Founda- 
tion). A grant from the National Science Foundation to the Population and 
Evolutionary Biology Program at Washington University in support of graduate 
training is also gratefully acknowledged. My thanks to Dr. John McNeill 
and an anonymous reviewer for helping to clarify the manuscript. 

280 Sida18(1) 


VoucluT specimens aiul lierbariiim of\)n^nn (ueronyms Follow Holmi;ren et til. 1990); chose lack- 
ing herbarium designation are housed at MO. For new combinations and new species in Leptucbluct see 
Snow 199Ha, b and Snow and Simon 1997. 

LeptoMoci .iqihUiai Scribn. & Merr.: Hitchmk 7()<)4 (US); Pr/rii^/e 6664 (US); .Vw/cnYrow 650 (US); MiVa//}^/} 

19124 a^S):Siioir662_'i. 
L. caiuiata (K. Schum.) N. Snow: Van Somereri AH9575 (US); Smm-den 1429 (US); Bogclciii I jO (UC). 
L. chinensis (L.) Nees: Snow et al. 698()\ Kanm 254 (K); Poore 440 (K); Dariclse 7471 (K,MO); Chiytun 

5644 (K). 
L. chlondifonius (Hack, ex Stuck.) Parodi: Biniheu HO (LP); S//re//s 622 (MICH); Peckrwi 3471 (US); 

PeJersefi 2662 (US). 
L. coerideueus Sreud.: Adiiw ni75\ At/am 14(>3(>:. Ad/w 5694. 
L. decipiens (R. Br.) Stapf ex Maiden subsp. cnthetms (Roem. & Scluilt.) N. Snow: Swiw & Simon 7272; 

Snow & Simon 732^; Snow & Simon 7335\ Cnsp et al. 2710 (MEL); 'rimw/Kuni & Slh/r/iv HUGin 

(BRI); Hiddhifd 509S(K). 
L. decipieus subsp. decipiens: Snow et ul. 724'^; Snow & Simon 732bl; Snow & Snnon 7334; B/ake 22548 

(CANB); Roes.n. (MEL, accession 234696); Ljz.nides 5634 (US); Rei^an .un. (CANB). 
/.. deap/en.\ subsp. /'tv/c«i// (Maiden & Betclie) N, Snow: S)iow & Simon ^ 323\Siiow & Simon 7329; Snow 

& Simon 7330; Snow & Simon '^336; Pi/rdie 3 1 5D (liR\); Boonnan .i.n. (G, accession 8227-(S6); /w/jw.uw 

713 (CANB). 
/,. digitataiR. Br.) Domur. Snou et ,d. 7224;Snowef al. 7 235;Siiow et al. ^246; B//rhidge 5326 {CAbiW); 

Blake 1 1 506 (CANB); Blake 6320 (CANB); Walter & Walter 2590 (B). 
L. divaricatisiima S. T. Blake: Snow et al. 7228; Snow et al. ^2)' i; Snow et al. 7236; Snow et al. 7241; 

Lloyd y^9 (CANB); Blake 77^7 (BRI). 
L. dnbia (Kuni\\)neiis: Snow 5865; Warnock46783 {NCU); Krai 5 1 801; Mearm 1213 (US); Hernandez 

& Mathns N-2066 (GH); Gould 12183 (K); Castillon 4356M (GH). 
L, eleiisme (Nees) T. A. Cope & N. Snow: Snow et al. 6941; Snow & Burgoyi/e 6)954; Snow & Biirgoyne 

6963; Snow et al. 6982; Schweiekerdt 1896 (PRE); Gi/y & Ward ^ (PRE); Drlge s.n. (S, accession 93/ 

1 94); E.xtenuon 0/fwer 164 1 9 (PRE). 
L. /n.\-ea (L.) Kiinth subsp. ///.kv/: Snow et al. 7215; S)iow et al. 7216; Snow et al. 7 222; Snow et al. 7232; 

Snow et al. 7234; Snow et al. 723^; Tracy 929? (GH); Pry s.n. (GH). 
L. fnsca s.u6-s\\. fascicidaris (Lam.) N. Snow: Snow 5786A; Snow 5800; Snow 5804; Snow 5809H; Snow 

581 ! A; Snow & Koster 5824; Snow & Koster 5840; Snow 584 1 ; Snow 5842; Snow 5896; Snou 5900; 

Snow 5901 -B; Snow 590 1 -H; Snow 5903; Praser 63 I (MICH); Pringle 9595 (MEXU); Swallen 4265 

(BAA); Wooton s.n. (BAA). 
/,. Ji/sca subsp. mnelleri (Bcnth.) N. Snow: Patz 1 1 (BRI); Maconochie 13433 (BRI); Ab/st 48^ (CANB). 
L. f/isca subsp. unmervia (J. Presl) N. Snow: Snow & Prinzie 648-i;Snou- & Priiizie 6567; Snow & Prinzie 

6568; Snow 6598. 
L. gigantea (Eaunert) T. A. Cope & N. Snow: Smi/li 4 1 26 (PRE); Smith 13H7 (BRI). 
L. ligulata Lazarides: Snow & Simon 7324; Snow 7402; Story & Yapp 25 (CANB). 
L. longa Griseb.: Davidse 2612; Soderstrom 10^3 (US); Hitchcock 10377 (US). 
L. malayana (C. E. Hubb.)Jansen ex Veldkamp: Yacoh 25883 (P). 
L. marqtiisensis (R Br.) P. M. Peterson & Judw.: Perlman 14919; Perlman 15064. 
L. monticola C.lvAst: Holdridge 1955 (US); Pkman HI 1874 (US). 
L. nealleyi Vasey: Snoir 5793B; Snow 5H05A; Snow 5920; Cory 20298 (GH); Waller & Banml 3096 

(GH);Johnston 4872 (MK:H). 
L. neesii (Thwaires) Bench.: Snow & Simon ^378; Snotc & Simon 7 384; Davidse & Siimithraarachcht 9180 

(MO.US); White 8894 (US). 
L. obtiisiftora Hochsc: Milne-lledhead & Taylor 7297 (B); Greenway 10666 
L. panicea (Recz.) Ohwi subsp. panicea: Reekmans 4982 (B); Ahmad s.n. (B, accesssion 173193-120); 

Hitchcock 194^4 (US); I'leld & Loew 6B (US); Backer 35094 (US). 
L. panicea subsp. brachiata (Sceud.) N. Snow: Snow 5905;Snow 5910A;Snow & Prinzie 6529C; Snow & 

Prinzie 6558; Snoir 6654; Snow 667 2-A . 
L. panicea subsp. nuicronata (Michx.) Nowack: Siiow 5847 A 

Snow, Caryopsis morphology of Leptochloa 281 

L paniandei (J. Presl) Hicchc: Snmv ':,192A\Sn„w 5SI()A; Snow 6622\Jacob 511 (NCU); Fnimoer & 

Williams 41 (ENCB); McDcimd & Rimaihi 18020. 
L. rupestris C. E. Hubb.: Wood 2848 (BM); Wood s.n. (K); Gilbert & Phillips 8874 (K). 
L. scabni Nets: Snow 5788; Snow 5791A; St/oir 5S/(M; McKenzie & Urhatsch 160 (NY); Nee 31921 

(NY); Burger & Ltesner 6958 (NY). 
L. southwoodii N. Snow & B. K. Simon: Snow & Stiiuni 7350; Snow & Simon 7362. 
L. squarrosa Pilg.: Greenway 2764 (P); Schliebtn 6>940 (P); Milne-Redhead & Taylor 7301 (B). 
L. srilankensis N. Snow: Davidse & Siniiithraaracbihi 9066 (K); fnsber^ et al. 50835 (CANB); Clayton 

5591 (TABS). 
L. tectoneticola {E&cker) ]dnsen ex Veldkamp: Poilam 15394 (P); Smitinand 3418 (CANB); Kerr 20633 (K). 
L. uniflora Hochst. in A. Rich.: Trimens 28 (US); Rains 67 (US); Thollon 4093 (US). 
L. virgata (L.) P Beauv.: Pohl & Davidse 10603 (UC); Burkart 22139 (SI); Meza 12; Swallen 3109 (US); 

Brockmann 18660 (BAA). 
L. viscida (Scribn.) Beal: Snow 6597; Riii'z s.n. (PNCB); Palmer 1 789 (CJH); Goodding & Lusher 139-45 

L. xerophila P. M. Peterson & Judw.: Halle 2098 (P). 
Cypholepis yemenciais (SchweinF.) Chiov.: Bally & Melville 1 5783; Verdcourt 3275. 


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National Herbarium of New South Wales 

Royal Botanic Gardens 

Sydney, N.S.W. 2000, AUSTRALIA 


Herbarium, Institute of Biology 

University of the Philippines 

Dilvnan 1101. Quezon City. PHILIPPINES 


A new species oi'Xdnthostemofi. X.fri/ticosus. presently known only from the Palanan Wilderness 
in Isabela Province, Luzon, is described and its relationships discussed. 


Una nueva especie de Xarithostemon, X. fruticosus, en la actualidad solamente conocida de 
la Palanan Wilderness en la provincia de Isabela es descrita y sus relaciones con otras especies 

Xanthostemon is a genus of around 45 species that occurs in Australia, New 
Caledonia, the Solomon Islands, Papua New Guinea, Indonesia (Irian Jaya, 
the Moluccas and Sulawesi) and the Philippines. In the Philippines, only 
four species have been named and there has been little published on the 
taxonomy of the genus since Merrill (1952) clarified the application of the 
names X, speciosus and X. verdugonianus. 

In 1991, a preliminary inventory of the Palanan Wilderness was under- 
taken as a joint project of Conservation International, Leiden University 
and the Isabela State University. A report of this inventory is given by Co 
and Tan (1992). One of the vegetation types encountered was a low scrub 
that occurred on ultrabasic substrate and was dominated by a shrubby 
Xanthostemon species with large, right red flowers. This has proven to be an 
undescribed species, most closely related to X. verdugonianus and is here described 
as X. fruticosus. 

Xanthostemon fruticosus Peter G. Wilson & L. Co, sp. nov. (Fig. 1). Type: 
PHILIPPINES. Luzon Island. Isabela Prov.: Aubarede Peninsula, Lanay Spring, ca. 
17 km NNW of Palanan Point, 25 May \39\,Uo)utrdoCo '^5S5 (holotype: A; isotypes: 

SiDA 18(1): 283-286. 1998 


SiDA 18(1) 

Fk;. 1 . XiinthdStciiiiJii jr/niius/is Peter Ci. Wilson & L. Co. A. Habir B. Immature fruit show- 
ing neatly circumscissile hypanthium. (;. Open, mature truit. D. Detail of leat showing 
venation. All from holotype. Scale bar: A, D=.7 cm; B, C=l cm. 

Wilson and Co, A new Xanthostemon species 285 

Xanthostemon verdugiano hypanthio circumscisso valde affinis sed habitu friiricoso foliis 
floribusque maioribus pedicellis pedunculisque longioribus et petalis roseis diftert. 

Shrub ca. 1 m high; young shoots sparsely appressed-hairy, glabrescent. 
Leaves spirally arranged; petiole 7-10 mm long, 2.5-3 mm wide; lamina 
obovate, 6-8 cm long, 3.5-4 cm wide, length:breadth ratio 1.6-2.1:1, co- 
riaceous, apex rounded and emarginate; oil glands numerous. Inflorescences 
crowded toward the apex of seasonal growth unit, subtended by slightly 
reduced foliage leaves, axillary, the apical bud apparently always growing 
on (i.e. shoot auxotelic). Unit inflorescences triads; peduncles 5-9 mm long, 
the upper ones shorter giving a corymb-like appearance to the flowering 
shoot; pedicels 6.5-10 mm long. Hypanthium cup-shaped, 5-6 mm in diameter, 
4—5 mm deep, exceeding the summit of the ovary, ±glabrous. Petals 5, pink, 
oblong to obovate, 5.5-8 mm long, 3.5-6.8 mm wide. Sepals 5, +equal, 2-2.7 
mm long, 2.5-3.8 mm wide, ±triangular, apex obtuse to rounded. Stamens 
red ±equally spaced, ca. 22 in a single series around the rim of the hypanthium; 
filaments 22-25 mm long; anthers 1.2-1.5 mm long, connective broad with 
one large gland at the apex and 2-5 other, smaller glands. Ovary partly in- 
ferior, (2-)3-locular, glabrous. Style 30-35 mm long, extending 3-5 mm 
beyond the stamens; stigma as wide as or slightly narrower than the style, 
flat. Fn,?i/ depressed-globular, ca. 8 mm diameter, ca. 6.5 mm high, broadly 
lobed at each loculus; hypanthium neatly circumscissile 1.5—2 mm above 
the base, the remnant ±flattened under the capsule. Placentas rod-like, horizontal. 
Seeds not seen. 

Distribution. — Known only from the type locality. 

Ecology. — Xanthostemon friittcosNS has been found near the coast in a low 
scrub community occuring on exposed sites on soils derived from an ultra- 
basic substrate. It is the dominant component of this community type at 
Lanay which also includes a number of other genera of spreadiing or erect 
shrubs (Co & Tan 1992). 

Conservation status. — ^Apparently rare. Only recorded from one of four ultrabasic 
sites visited in this part of north-eastern Luzon. Ultrabasic areas are very 
marginal for agriculture but are potentially subject to applications for nickel 
mining. As a wilderness, the region does have legal status as a 'Protected 
Area' but this may not be an impediment to mining. 

Etymology. — The epithet is derived from the 'L2itm,fruticosus, shrubby, referring 
to the distinctive habit, a characteristic that distinguishes it from the re- 
lated X. verdugonianus . 

The red-flowered X. speciosus Merr. From Palawan and nearby islands has 
a broader, dish-shaped hypanthium with distinct vesicles and is thus not 
closely related to the new species but belongs with X. confertiflorus (Sulawesi) 
and X. youngii (north-eastern Australia) in sect. Vesicaria of Gugerli (1940). 

286 SiDA 18(1) 

Xanthoste?nun speciosus is usually a tree of 5 meters or more but small trees or 
shrubs of only 1 meter or so in height have been recorded (e.g. Co 3032, A, 

Xanthostemon fniticosus is closely related to X, verdugonianus which it re- 
sembles in having red flowers with a circumscissile hypanthium. Xanthostemon 
verdugonianus is a tree up to 26 m high that ranges from north-eastern iVIindanao 
to Sibuyan Island, the latter being over 500 km south of the Palanan area. 
Despite its range, X. verdugonianus has a very limited habitat and is much 
sought-after for its durable timber, making it a threatened species (Yao & 
Ulep 1981, 1983). 

The neatly circumscissile hypanthium appears to be unique to this pair 
of species. Two species in New Caledonia have hypanthia that split and are 
irregularly circumscissile but these are members of an endemic group of 
taxa with buUate leaves (sect. Bullata of Gugerli 1940) and do not appear 
to be closely related to the Philippine species. 

The species may be distinguished as follows: 

1. Shrub ca. 1 m tall; lamina 6-8 cm long, 3.5-4 cm wide; petiole 7-10 mm 

long; petals pink X. fruticosus 

1. Tree to 26 m tali; lamma 3-7 cm long, 1-3.5 mm; petiole 4-7 mm long; 

petals red X. verdugonianus 


The first author is grateful to the directors of A and CANB for loans of, or 
access to, herbarium specimens examined for this paper. Thanks, also, to 
Lesley Elkan for the illustration and Jaime Plaza for assistance with the Spanish 
abstract. The second author thanks Conservation International for funding 
his field work in the Palanan Wilderness in 1991- 


Co, L.L. and B.C. Tan. 1992. Botanical exploration in Palanan Wilderness, Isabela Prov- 
ince, the Philippines: first report. Fl. Males. Bull. 1 1 :49-53. 

Gugerli, K. 1940. Monographic der Myrtaceengattung Xanthostemon. Reperc. Spec. Nov. 
Regni Veg. Beih. 120:1-149. 

Merrfu, E.D. 1952. Notes on Xanthostemon F. Mueller and Kjellbergiodmdnm Burret. J. Arnold 
Arbor. 33:150-165. 

Yao, C.E. and E.V. Ulep. 1981, Mangkono in Babatngon. Canopy 7:5. 

Yao, C.E. and E.V. Ulep. 1983. More on Mangkono. Canopy 9:6-7. 









Laboratory for Environmental Biology and 

Department of Biological Sciences (WPM RM, CEF, HN.JG, HN, KR) 

Department of Geological Sciences (NEP) 

University of Texas, El Paso, TX 79968, U.S.A. 


We compared seasonal changes in concentrations of four elements, arsenic, cadmium, 
copper and lead, in samples of various tissues of creosotebush collected in the El Paso, USA/ 
Ciudadjuarez, Mexico region during 1980-81 and 1994-95. Levels in leaf tissue changed 
seasonally, with concentrations dropping in the spring and late fall, corresponding to the 
time of leaf drop in the plant. This suggests that most of the heavy metals were simply 
deposited on the surface of leaves, although data are presented which indicate that internal 
tissues also have significant levels of heavy metals. These seasonal cycles were less pronounced 
in the most recent samples. Levels of all four elements in cresotebush were significantly 
high in the region, as compared to a distant control area. Concentrations of most elements 
dropped below detection limits within 12 to 30 km from areas of highest concentration. 
Densities of native Chihuahuan Desert flora and lichens are low in the area, suggesting an 
apparent negative impact of industrial pollution on the local vegetation. Cadmium and 
lead levels in creosotebush tissues have dropped over the past 15 years, suggesting that 
enforcement and strengthening of environmental laws has reduced the air pollution levels 
in the El Paso area. 


Comparamos los cambios esracionales de las concentraciones de 4 elementos, arsenico, 
cadmio, cobre y plomo, en muestras de varios tejidos de gobernadora colectados en el area 
deElPasoy Ciudadjuarez, Mexico durante 1980-81 y 1994-95. Se encontro un cambio 
estacional de los metales con las concentraciones disminuyendo en la primavera y otono, 
que coresponden al momento de la caida de las hojas de la planta. Esto sugiere que hay 
deposicion de los elementos en la superficie de las hojas, aunque presentamos datos de que 
hay acumulacion de metales en los tejidos interiores. Estos ciclos estacionales estan menos 
pronunciados en los ultimos anos. Los niveles de los elementos son altos en la region, pero 
estan concentrados en un area entre 12 y 30 kilometros del area con concentraciones mas 
altas. Las densidades de la flora y Ifquenes en el area son bajas, suginendo un impacto negativo 
de lapolucion industrial en la vegetacion local. Los niveles de cadmio y plomo en los tejidos 

SiDA 18(1): 287-296. 1998 

-«8 SiDA 18(1) 

ha disminuiclo en los ultimos 1 5 anos, lo que siii^iere que la aplicacion cle las leyes medioambientales 
ha reclucido los niveles de polucion en el area de El Paso. 

The El Paso/Cd. Juarez area is one of the many regions along the United 
States/Mexico border which suffers from environmental contamination. The 
area has been subjected to heavy industrial activity, including smelting and 
hydrocarbon refining, for over 100 years. This has resulted in an accumula- 
tion of heavy metals, especially lead, cadmium, copper and zinc in the local 
soils (Barnes 1993; Ndame 1993). Arsenic contamination has been previ- 
otisly documented to occur in this region of west Texas (Shields 1991). 

These elements and their compounds can cause medical problems in humans 
and other animals (Elbahri & Benromdane 1991 ; Louekari et al. 1991). Negative 
effects of heavy metals on plants have been documented (D'ltri 1982; Fernandes 
& Henriques 1991), including the prevention of the uptake of potassium 
and phosphorous by roots. Moreover, copper may damage chlorophyll and 
increase the potency of fungal diseases (Connell & Miller 1984). These ef- 
fects may eliminate some plant species, with concomitant increases in the 
abundances of others, thus changing plant community structure. Plants may 
tolerate heavy metal contamination (Connell & Miller 1 984) by excluding 
metals from sensitive tissues, modifying metabolic pathways to prevent damage 
or assembling specific enzymes to detoxify heavy metals. The specific ef- 
fects of heavy metals on the flora of the northern Chihuahuan Desert have 
not been documented, but Worthington (1989) previously reported a re- 
duction in species richness in native plants in this area, presumably the result 
of such contamination. 

In this investigation, tissues of the dominant Chihuahuan Desert shrub, 
Larrea tridentata were analyzed, to document continued heavy metal con- 
tamination in western Texas. This work is part of a larger study on the ef- 
fects of heavy metal contamination on the flora and fauna of the northern 
Chihuahuan Desert and the investigation of bioaccumulation into higher 
trophic levels. 


We selected a site on the campus of the University of Texas (UTEP) in 
western El Paso to follow seasonal change in heavy metal concentrations m 
creosotebush. The site is located within 2 km of a local smelter, which ap- 
pears to be a point source for the high concentrations of heavy metals in the 
area. Four elements were included in the analysis: Arsenic, Cadmium, Copper 
and Lead. Additional sites used to determine the distribution of heavy metals 
in creosotebush tissues in the region included 62 sampling areas near the 
border in the United States and six in Mexico within 10 km of the border 
and ot the smelter, of which 52 yielded creosotebush specimens (five repli- 

Mackay, et al., Heavy metals in Larrea cridencaca 289 

cate bushes at each site). Sites were selected which had native Chihuahuan 
Desert vegetation located at least 50 meters away from any road. Prelimi- 
nary analyses showed higher levels ot lead next to roadways, which decreased 
to background levels within a few meters, an effect also noted by Motto et 
al. (1970), Gratani et al. (1992) and Lebreton and Thevenot (1992). 

Leaves were stripped, and twigs were cut from the plants. Roots and trunk 
tissues were also harvested from plants. Tissues were placed in numbered 
paper bags and returned to the laboratory. The samples were further cleaned, 
removing all foreign matter, but were not washed. Bark and exterior tis- 
sues were carefully removed from pieces of trunks, to avoid contamination 
of internal tissues. Tissues from 1980—1981 were collected North of the 
Education Building on the UTEP campus, as part of another study (Free- 
man 1982). The tissues were ground and stored dry in glass vials until they 
were analyzed in 1994. Tissues from 1994—1995 were collected through- 
out the season from five specific bushes, located in Charlie Davis Park, on 
the campus of UTEP. This allowed a comparison of the percentages of heavy 
metals in the tissues of each of the five creosote bushes. 

Approximately 20 grams of tissue were placed in a crucible and muffled 
for three days at 425°C. This relatively low temperature was selected to 
avoid vaporizing the four elements or their salts (based on the recommen- 
dations of Dr. Jim Rayon and Dr. Ken Dodson of the Environmental Pro- 
tection Agency). After ashing, 100 mg of ash were dissolved in 20 ml of 
14% nitric acid (V:V), without filtering. Reagent grade acid and double 
glass distilled water were used for all solutions. Samples were prepared and 
stored in glass scintillation vials with polypropylene-lined caps. 

Samples were analyzed on a Beckman SpectraSpan 6 direct current plasma 
atomic emission spectrograph (DCP-AES). We followed the protocol of EPA 
method 6010 for the inductively coupled plasma emission spectrometer (US 
EPA 1986). Three of the wavelengths used are those recommended by EPA: 
193-696 nm for arsenic, 213.598 nm for copper, and 220.353 nm for lead. 
The fourth wavelength, 228.802 nm, was substituted for the cadmium analysis 
to achieve the lowest detection limit. The DCP was calibrated with pre- 
pared standard solutions such that the linear calibration curve had an R- of 
at least 0.995. The samples were analyzed by DCP three times and reported 
the mean and standard deviation of these readings. If the relative standard 
deviation was greater than 3%, the sample was reanalyzed. Blanks, dupli- 
cate samples and prepared standards (spiked samples) were analyzed at least 
once every ten samples for quality assurance/quality control. Cadmium 
concentrations in the samples collected in 1994—1995 were close to or be- 
low the limits of detection and therefore we have not presented them. Af- 
ter analysis, the DCP solution values were converted to ppm in the original 

290 SiDA 18(1) 

ash. The detection Hmits in the ash for this study were 200 parts per mil- 
Hon (ppm) for arsenic, 11 ppm for cadmium, 85 ppm for copper and 130 
ppm for lead. 

Data were j^lotted with SURFER (Golden Software Inc., Golden Colo- 
rado), using the grid method with inverse squared distances and the sur- 
fdce module (Keckler 1995). 


Seasonal changi^s. — Levels of all four elements varied seasonally during both 
time periods (Fig. 1). Levels were high in the winter and showed a small 
drop in the sj^ring when there is a combination of minor leaf drop and strong 
spring winds. The levels increased during the summer, especially during 
the early sampling period (Figs. 1 a, b, c), but later showed a large drop during 
the fall when creosote lose a large proportion of their leaves (Mackay et al. 
I 987). (Concentrations increased again during the winter months, especially 
during 1980-1981 . Levels of heavy metals were lower in the 1994-1995 
samples (Fig. Id & f). Cadmium and lead were both approximately 4 times 
higher in 1980-1981, suggesting that enforcement and strengthening of 
environmental laws has reduced the air pollution levels in the El Paso area. 

Seasonal changes in concentrations of all metals during both sampling 
periods were statistically significant (Table 1). The first harmonics from Fourier 
analysis of the means (Little & Hills 1972), corresponding to the seasonal 
effect, were significant for all years, although the patterns were more pro- 
nounced during the first sampling period (Fig. 1) and had correspondingly 
higher F values (Table 1). Other harmonics were statistically significant in 
1980-198] , showing the importance of leaf drop in lowering heavy metal 
concentrations during these years, although there was no pattern in which 
harmonic was significant after the first. The lack of significance of higher 
harmonics during the second sampling period suggests that leaf drop was 
not significant in reducing heavy metal content during those years. 

Contani'iucttion of tissues. — It appears that large amounts of the heavy metals 
arsenic, copper and lead are deposited on the leaves, but heavy metals are 
also incorporated into the tissues of this plant (Fig. 2). Large concentra- 
tions of all three elements were found in the roots. The trunks also had high 
concentrations of the three elements, which were present in the internal 
tissues as well. The branches and the leaves also had high concentrations of 
the three elements. These data suggest that heavy metals may be deposited 
on the outside of the plant, but is also taken up by the roots and incorpo- 
rated into the woody tissue. 

Heavy metal geographical distribution: Levels of the four elements (ar- 
senic, cadmium, copper and lead) in creosotebush ash were elevated adja- 
cent to the LJnited States/Mexico border on the west side of El Paso (Fig. 


;kay, I 



















T Al,., Heavy metals in Larrea tndentata 



400 - 



300 - 



-p T 

200 - 




100 - 




'^ J_ I? 



1 1 1 I 

^ d) 

1 1 



T 1994 - 199f 


5000 ^ 



4000 - 

I 1 

3000 - 


k T < 

1 1 




2000 - 
1000 - 




•— ir* 


r ^1 



1 1 

Pb c) 

n \ 1 1 1 1 r 





O D 



Fir,. I: Seasonal changes in heavy metal concentrations in the leaves of the creosotebiish, 
Larrea tridentata dnrmg 1980-1981 and 1994-1995. The gap in 1994-1995 is due to 
lost samples. Error bars represent standard error of the mean, and are not included in a, b, 
and c as many ol the data points are based on a single sample. The months are abbreviated 
by a single letter on the x axis. 

3). Levels of cadmium were as high as 190 ppm in ash, chose of copper reached 
5200 ppm and lead levels were as high as 1200 ppm (Fig. 3, note that the 
surfaces in the figures are close to average values and do not extend to these 
extreme values). The highest levels of the three elements were recorded on 
the east side of Mount Cristo Rey {Peak 1 in Fig. 3a), on a small mesa west 
of McNutt Road (Peak 2 in Fig. 3a), and on the UTEP campus (Peak 3 in 
Fig. 3a). Levels of all three elements were lower on the southwest side of 
Mount Cristo Rey, perhaps due to a wind-shadow effect. Levels of all three 
elements rapidly decreased to the east and fell below detection limits at 
distances of between 12 and 30 km (Fig. 3). 


SlDA 18(1) 

Tabu; 1 . Resulrs oFFourier Analysis (Little & Hills 1972) of'tht' periodic tunctions in Figure 1 . Meuns 
were used m the analysis. 


df Mean Squares 

Arsenic, ] 980-198 I 

F Values 

First harmonic 

Second harmonic 

Third harmonic 

Fourth harmonic 

Fifth harmonic 




2.2 ns 
3.7 ns 
0.9 ns 

Cadmium, 1980-1981 

First harmonic 

Second harmonic 

Third harmonic 

Fourth harmonic 

Filth harmonic 


I'lrst harmonic 

Second harmonic 

Third harmonic 

Foiu'th harmonic 

Filth harmonic 





















Lead, 1980-1981 






180191 1.0 















1.9 ns 


IC) C)*** 

0.8 ns 
1.1 ns 

3.7 ns 

Arsenic, 1994-1995 

First harmonic 

Second harmonic 

Third harmonic 

Fourth harmonic 

Fifth harmonic 






1 159.2 
1 123.4 
61 1 1.7 


0.3 ns 

1.7 ns 
0.2 ns 
3. 1 ns 

Mackay, 1-t al., Heavy metals in Larrea tridencata 293 

Tabu: I . continued. 

Source df Mean Squares F Values 

Arsenic, 19.S0-I981 

Copper, 199i 


I'irst liarmonic 


1 168398.7 





Second harmonic 



1.8 ns 




Third harmonic 



1.2 ns 




Fourth liarmonic 


1 18186.9 

0.5 ns 




Fiftii harmonic 



1.7 ns 



Lead, 1994- 


First harmonic 







Second harmonic 



0.9 ns 




Tiiird liarmonic 



0.8 ns 




I'ourth harmonic 



0.1 ns 




Fiftli harmonic 



0.9 ns 




* = F value of harmonic 

sit,'nificant ; 

ut 0.05 level, ** 

= F significant at 0.01 level 

, *** = Fsignifi- 

cant at ().()() 1 level, ns - 

F" vaJLie not 


The areas with the highest values of As, Cu, Cd, and Pb in creosotebush 
(peaks 1-3, Fig. 3) coincide with the sites of the highest concentrations of 
heavy metals in the soils (Barnes 1 993; Ndame 1 993) and in fluff grass (MacKay 
et al. 1998). In this locale, Pb in the soils exceeds the EPA TCLP (Toxicity 
Characteristics Leaching Procedures) regulatory limit at a number of sites, 
and Cd is reported quite close to the limit. The spatial correspondence between 
elevated metal levels in the soil and the flora is not unexpected. The mechanism 
of uptake of heavy metals by creosotebush remains to be elucidated. 

Densities of native Chihuahuan Desert flora and lichens are low in the 
area (Worthington 1989, MacKay et al. in prep.), suggesting an apparent 
negative impact of industrial pollution on the local vegetation. The spe- 
cific effects of heavy metals on the flora of the northern Chihuahuan Desert 
are currently being documented. The demonstrated reduction in species richness 
in native plants in this area is presumably the result of heavy metal con- 
tamination from the smelter. These effects may eliminate some species and 
increase the abundance of others. 


SiOA 18(1) 

I I Roots 

y//////A Trunk 

^^ Branches 

Nxxxxxx?! Leaves 

I I Bare Trunk 

Fic,. 2: The- tlisrnbutions of lead, arsenic and copper in specific tissues of the creosotebush 
from Ciiarlie Davis park on the University of Texas, El Paso campus. El Paso, TX. 


We would like to thank the United States Department of Agriculture, 
especially J. A. Vigil and J. Underwood, for permission to import plant tis- 
sues from Mexico. The Texas Department of Parks and Wildlife, especially 
David Riskind, allowed tis to collect plant tissues in the Franklin Moun- 
tains State Park. Ken Dodson and Jim Rayon offered suggestions regard- 
ing sample preparation. Our research was supported by the Center for 
InterAmerican and Border Studies of the University of Texas and by the 
National Science Foundation (IIRD 9253021). 


Barnes, B. 199.t. An evaluation of metals concentrations in siirficial soils. El PasoCotmty, 
Texas. Unpublished Master's Thesis, University of Texas at El Paso. 

Mackay, et Ai., Heavy metals in Larrea triclentata 




y. ^-C*** 






Fig. 3: The distribution of arsenic, cadmium, c()|")per and lead in the Ei Paso/Ciudad Juarez 
area. Charley Davis Park on the UTEP campus is located at the origin in the x-y coordi- 
nate system (0,0). Peak one corresponds to tiie east side of Mount Cristo Rey, Peak 2 to a 
mesa above (west of) McNutt Road and Peak 3 is on the UTEP campLis. Detection limits 
were 200 ppm for arsenic, 1 1 .2 ppm for cadmiimi, 84 ppm for copper and 132 ppm for 

CoNNF.i.i., D.W. and G.J. Miller. 1984. Cihemistry and ecotoxicology of polhition. Jolm 
Wiley and Sons, New York. 

D'llr], F.M. 1982. Acid precipitation: effects on ecological systems. Ann Arbor Science, 
Ann Arbor, MI. 

Ei.BAHKi, L. and S. Benromdane. 1991. Arsenic poisoning in livestock. Veterinary Human 
Toxicol. 33:259-264. 

Fernandes, J.C. and F.S. Henriques. 1991. Biochemical, physiological and structural ef- 
fects of excess copper in plants. Boc. Rev. 57:246—262. 

Freeman, C.E. 1982. Seasonal variation in leal nitrogen in creosotebLish {Larrea tridentata 
{DC.} Gov.: Zygophyllaceae). SouthW. JMatutalist 27:354-356. 

Gratani, L., S. Ta(;li()ni, and M.F. Crescen'it,. 1992. The accumulation of lead in agricul- 
tural soil and vegetation along a highway. Chemosphere 24:941—949. 

Keckler, D. 1995. Surfer for Windows, Ver 6.01, Golden Software, Inc., Golden Colo- 

Lebreton, L. and D.R. Tiievenot. 1992 Polkition metallique relargable par les aerosols 
d'origine autoroutiere. Environ. Technol. 13:35-44. 

LrnxE, T.M. and F. Hills. 1972. Statistical methods in agriculrural research. Agric. Ext. 
Service, Univ. California. 

296 SiDA 18(1) 

LouHKAR), K., S. Valkonhn, S. Pcmsi, and L. Virtani;n. 1991 . Estimated dietary intake of 
lead and cadmium and tlieir concentration in blood. Sci. Total Envnx)n. 105:87—99. 

iViACKAY, W.F'., S. Silva, S. Lorinc, and W. Whitford. 1987. The role of subterranean ter- 
mites in the decomposition of above ground creosotebtish litter. Sociobiohjgy 1 3:235— 

JVIackay, W.P., L. l^E La Rosa, M. Zavala, P. Suhton, J. Gari)[;a, R. Mi;na, N. Pinc;ii()ri;, 
and G. C^RAWfORD. 1998. The distribution ot heavy metal contamination m the El Paso, 
TX/Ciudad Juarez, Mexico area, usmg fluff grass, Er/one/in))i juilchelli/rti, as an indicator 
organism. Proc. Chih. Desert Res. Inst. (In Press). 

Morw) I I.E., R.H. Dainhs, D.M. Ciiii.ko, and C.K. Mcrrro. 1970. bead in soils and plants: 
its relationship to tralfic volume and proximity to highways. Environ. Sci. Technol. 4:231 — 

Ndamb, E.G. 1993. Heavy metals in soils in the vicinity of the University ol Texas at El 
Paso campus (El Paso, Texas). LJnpubl. Master's Thesis, LJniversity of Texas, El Paso. 

Shields, J. 199 1 . Ambient air arsenic levels along the Texas - Mexico border. J. Air Waste 
Manag. Assoc. 41:827-831. 

LIS Environmental Proiec-iion Ac;ency. 1986. Test methods for evaluating solid waste 
(third edition). Washington, D.C. 

WoKiiiiNCiTON, R.D. 1989. Effects of El Paso pollutants on the lichen, moss, and winter 
annual flora on andesite rock formations. In: P. Ganster and H. Walter, eds. Environ- 
mental hazards and bioresource management in the United States/Mexico borderlands, 
UCLA Latin American Center Publications. Los Angeles. Pp. 105 — 115. 



Center jor B wdwer.uty 
Illinois Natural History Survey 

607 E. Peabody Drive 
Champaign. IL 61820. U.S.A. 


Division of Forestry and Wildlife 

Botanic Gardens 



Twenty-six vascular plant species are reported here for the flora of Dominica, in the Lesser 
Antilles. These include two pteridophytes, 13 monocotyledons, and 11 dicotyledons. Four 
of these species arc reported as new to the Lesser Antilles: Mar.ulea polycarjm Hook. & Grev., 
Dipldcriim Ccipttciliini (WiUd.) Boeck., Rhy)ichoipi)rci niceiiiosn Wright ex Sauvalle, and Phariis 
lLlppnlclCt!lS Aubl. 


Se citan 26 plantas vasculares adicionales para la flora de Dominica en las Antillas JMenores. 
Estas especies inckiyen dos pteridolitos, 1 3 monocotiledoncas y 1 1 dicotiledoneas. Cuatro 
de estas especies se presentan como nncvas para la flora de las Antillas iVIenorcs: iWarsilea 
polycarpa Hook. & Grev., Diplacr//iii aipiu/i/nii (Willd.) Boeck., Rhyrichospora racemout Wright 
ex Sauvalle, y Vloarin lappulaceus Aubl. 


Dominica is a volcanic island located between the French islands of Guadeloupe 
and Martinique in the Lesser Antilles. It is about 45 km long and 24 km 
wide. While it is a small island (1088 sq. km), it presents a diversity of 
habitats along altitudinal and moisture gradients because of its rugged 
mountainous topography. Several of the mountain peaks exceed 1200 m 
(MorneDiablotins 1433 m, Morne Trois Pitons 1402 m, Morne Watt 1242 
m, Morne Anglais 1223 m). Water is abundant on the upper slopes of the 
mountains (e.g. at Freshwater Lake, 8459 mm of rain/yr. has been recorded) 
but rainfall on the western (Caribbean) coast is severely restricted by a rain 
shadow (< 2000 mm/yr.) and a dry scrub forest prevails. Severe damage caused 
by hurricanes and agriculture has also provided opportunities for opportu- 
nistic species, increasing the diversity. The coastline is very steep, and the 

SiDA 18(1): 297-305. 1998 

298 Sida18(I) 

cliffs generally continue precipitously into the ocean. Level land is prima- 
rily restricted to river deltas in a few narrow bands near the coast, where 
most citizens live. 

Botanically, Dominica is rather well-known. The dicotyledonous flora 
was treated by Nicolson ( 1 99 1 ) who estimated a vascular flora of 1226 spe- 
cies. The most recent treatment of its pteridophytes and monocots Wiis Howard's 
?,\x\o[ivm(^ Flora of the Lesser Anttlles {\91 A, 1977, 1979, 1988, 1989a, 1989b) 
which included detailed distribution notes within that region, and treated 
the dicots as well. Whitefoord (1989) added 40 phanerogams and six ferns 
to the flora. This paper reports twenty-six vascular plant species newly dis- 
covered and vouchered for Dominica. 


The plants reported here as new to Dominica were discovered during the 
period 1990-1997. Approximately 2()()() collections were made by the first 
author lor the purpose of assembling a reference herbarium at the Spring- 
field Estate, formerly called the Archbold Tropical Research Center after 
the late John Archbold who donated the property to Clemson University 
in 1989. The Springfield Estate is currently leased from Clemson Univer- 
sity by a Dominican non-government organization, the Springfield Centre 
for Environmental Protection, Research, and Education (SCEPTRE). The 
herbarium there has been informally designated with the acronym "atrc." 
Specimens donated by several other collectors have also been incorporated 
into the herbarium. 

Collections (2005 numbers) have been made on Dominica during eight 
visits to the island by Hill as follows: 20 March-26 March 1990 (collec- 
tion numbers 27230-27.379), 4 March-23 March 1991 i22()()9-22]79), 
26july-5 August 1992 (23924-24171), 5 March-16 March 1993 (24612- 
24847, w/J. Gable & B. Dorsey), 7 December-1 9 December 1 993 (25309- 
23363, w/l.Renne & D.Bradshaw), 28 May-6 June 1994 (23610-23872), 
24 May-9 Jtme 1996 (27861-28262), and 23 Eebruary 1997-14 March 
1997 (28868-29109, w/L.R. Phillippe). Sets of specimens have been de- 
posited primarily in the Dominican herbarium (atrc), Clemson University 
(CLEMS), the Illinois Natural History Survey (ILLS), and the Smithsonian 
Institution (US). Additional duplicates have been distributed to herbaria 
cited after the individual specimens. 

New records of species generally considered to be indigenous to the Lesser 
Antilles (according to Howard 1974, 1977, 1979, 1988, 1989a, 1989b) 
are reported here. Some may have actually been introduced to Dominica by 
human activities, but direct evidence is lacking. 

Hill and James, Plant records for Dominica 299 




Trichomanei crinitum Sw. An epiphyte similar to Trkhomafies crispum L. but 
differing in its lobed or pinnatifid, not entire, pinnae. Its general distribu- 
tion is northern South America to Jamaica, and it has been reported from 
Guadeloupe, St. Vincent, and Grenada in the Lesser Antilles. We thank A.R. 
Smith for the identification. 

St. George Parish: elfin forest on windswept ridge between Breakfast River and Valley 
of Desolation, 6 Mar 1997, Hill & Phillippe 29070, (atrc, ILLS, UC). 


Marsilea polycarpa Hook. & Grev. A trailing perennial of temporary ponds, 
this species of northern South America has not been previously reported in 
the Lesser Antilles. It has several globose sporocarps arranged and attached 
in a linear fashion on the frond stipes rather than the single basal sporocarp 
seen in most other species of the genus. Neither the genus nor family have 
been previously reported from Dominica. 

St. Andrew Parish: locally abundant in and around temporary ponds at E end of Melville 
Hall Airport, Londonderry Bay, 23 Feb 1997, Hill & Phillippe 28868 (atrc, BRIT, ILLS, 
MO, NY, UC, VT). 




Agave dmstana Trel. The century plants are seldom collected, and none were 
reported from Dominica by Howard (1979)- This species, which is endemic 
to the Lesser Antilles, is very conspicuous in the dry scrub on the dry rocky 
hills of the NW coast of Dominica in areas of very low rainfall, growing 
mostly with cacti. Previous collections are known from St. Barts, Antigua, 
Montserrat, Guadeloupe, and Martinique. 

St. Joseph Parish: Morne Raquette (Rachette), Au Piton Road, 18 Dec 1993, H/7/ 2532-^ 
(atrc, NY). St. Peter Parish: Clement James' property, Anse a Liane trail to coast, 0.9 km 
N of Colihaut, 2 Mar 1997, Hill & Phillippe 28988 (atrc, BRIT, ILLS, MO, NY, TAES, 
USF, VT). 


Diplacr/iiii cctptlalnm (Willd.) Boeck. This species is rather common in Ven- 
ezuela and ranges from Panama to Bolivia as well. This appears to be the 
first record from the Lesser Antilles. Howard (1979) listed only Diplacrum 
longifoliiim (Griseb.) C.B. Clarke ex Dur. & Schinz. for the Lesser Antilles, 
from St. Paul Parish, Dominica. We thank JVL Strong for the identification. 

300 SlDA IS(l) 

St. Joseph Parish: Imperial Higiiway, Central I'Oresi Reserve, Dleu Gommier area, 6 
Jun 199 1, ///// JXSY}^ (acre, NY, US, USF). 

i'liNhristylis cymosci R. Br. subsp. spcithitceci (Roch) T. Koyama. This is a char- 
acteristic, wiry-leaved and toti^di-rooted, sedge of the windswept flats of 
the E and NE coasts of Dominica. Pantropical in cUstribution, it has been 
collected previously on most of the Lesser Antilles, including Gtiadelc:>upe 
and Martinitjue. 

St. Andrew Parish: roaclsiele 0.5 mi N ol Melville Hall Airport, Londonderry Estate, 
16 Mar 1993, ///// 24^42 (acre, CLEMS, MO, NY, TAES, USF). 

Rhyiich(is[>i>rc/ nwei/iosa Wright ex Sauvalle. This is the first report of this species 
for the Lesser Antilles. It is relatively common in the Greater Antilles. We 
thank M. Strong for the identificatit)n. 

St. Peter Parish: Morne Espagnol, Irom (xiasral Hii^iiway to stimmit, 4 Aiii,^ 1 992, ///// 
24/49 (CLEMS, NY). 


Brachiomdinvi parvnm Cogn. This tiny epiphyte of the upper montane rainforests 
was previously known from Venezuela and from Guadeloupe, Martinique, 
and Grenada in the Lesser Antilles. It was to be expected in Dominica but 
probably has been overlooked because of its small size (< 5 cm tall). We 
thank D. Nicolson for the identification. 

St. Peter Parish: NW side of Morne Oiahlotins peak, elfin forest, 6 Mar 1993, ///// 
24(^47 (ariT, CLEMS). St. George Parish: Lacidat, ea. I km W of Titou Gorge, 4 Jun I 996, 
///// 2HI77 (ati-e, ILLS, USE). 


khiianthns mnioroms (Sw.) Doell. van nvartzii K.E.Rogers. Generally distributed 
in the Antilles and Trinidad, this trailing rainforest species has been col- 
lected previously on St. Kitts, Nevis, Montserrat, iVlartinique, St. Vincent, 
and Grenada in the Lesser Antilles. 

St. George Parish: Morne Anglais trail to snmmir, S of Giracidel, 1 Jun 1996, Hill 
28124 (acre, ILLS). 

l.\achiiccii/i:^//.stif(jlui Nash. This tiiicket-forming grass which often climbs over 
other plants has been known previously from Puerto Rico, and from Guadeloupe, 
Martinique and St. Kitts in the Lesser Antilles. The new record is an ex- 
pected range extension. We thank G. Davidse for the identification. 

St. George Parish: Morne Anglais crail at siimmic, S ol Giraudel, 2 Jun 1996, ///// 
2.S'077 (acre, ILLS, TAES). 

Iscichne ctnnidnuiceci (Sw.) Griseb. This is another clambering grass of exposed 
wet mountain slopes, and its general range is Jamaica to Trinidad S to Bo- 
livia and N to Mexico. In the Lesser Antilles it has been previously col- 
lected on St. Kitts, Guadeloupe, and (irenada. We thank G. Davidse for 
the identification. 

Hill and James, Plant records for Dominica 301 

St. Paul Parish: trail to summit of Mornc Trois Pitons, N side, just NE of Pont Casse, 
9 Dec 1993, HjH 23340 (atrc, iVIO, VT). 

Leptochloa virgata (L.) Beauv. A very widely distributed grass which ranges 
from the southern United States through the Caribbean and Central America 
to South America. It has been collected on many of the Lesser Antilles, m- 
cluding nearby Marie Galante and Martinique, and was to be expected on 


St. Peter Parish: Clement James' property, Anse a Liane trail to coast, 0.9 km N of 
Colihaut, 26 Feb 1997, Htll & Phillippe 28962 (atrc, ILLS, MO, TAES). 

Pharus lappulaceus Aubl. This wide-ranging, wide-leaved grass is found in 

forests from the Caribbean and Central America to southern South America. 

Howard (1979) noted the similar?. glaberKvmth from Dominica. We thank 

T.S. Filgueiras for the identification. 

St. Mark Parish: Petit Coulibri,Morne Pons, 15 Mar 1993, H/7/ 24796 (atrc, CLEMS). 

Phragmites amtralh (Cav.) Trin. ex Steud. Probably due to the general lack 
of suitable habitat, the common reed has not previously been collected on 
Dominica. Otherwise, it is widespread in both tropical and temperate por- 
tions of the world, and has been collected previously in the Lesser Antilles 
on Guadeloupe, Martinique, and St. Lucia. 

St. John Parish: marsli at N side of Coconut Beach Hotel, Prince Rupert Bay, Ports- 
mouth, 29 May 1996, Hill 27993 (atrc, ILLS). 

Spartma patem (Ait.) Muhl. This grass, usually found in salt marshes and 
flats along the coast, was a surprising find in Dominica where these habi- 
tats are essentially lacking. The species generally ranges from the E coast of 
North America from Quebec to Mexico, Central America, and the Carib- 
bean. In the Lesser Antilles it has been collected previously on St. Martin, 
Antigua, and Guadeloupe. We thank D. Nicolson for the identification. 

St. Andrew Parish: just N of the mouth of the Londonderry (Tweed) River, Londonderry 
Estate, coastal rock ledges, 14 Dec 1993, Hill & James 2547 3 A (atrc, VT). 

Urochloaplantaginea (Link) Webster. Often reported as Brachtaria plantaginea 
(Link) Hitchc, this grass of unstable areas ranges widely from Mexico and 
the Caribbean to Brazil and Bolivia. As with Phragmites, it may be a recent 
introduction, though Dominica falls within its general range. It has been 
previously collected in the Lesser Antilles only on St. Lucia. We thank T.S. 
Filgueiras for the identification. 

St. Paul Parish: Morne Daniel Road, Canefield, 10 Mar 1993, 24701 (atrc, CLEMS). 



Conocarpus erecta L. The button mangrove (buttonwood) is widely distrib- 
uted in the American tropics along the coasts of Florida, USA, S through 

302 SiDA 18(1) 

Mexico and Central America ro the Caribbean and tropical South America 
and it has also been collected in West Africa. Buttonwood has been found 
on nearly all of the other Lesser Antilles. Hodge (1964) noted: "the com- 
mon genera of mangrove {Rh'izophora, Aviceimia, Lagunaihma, and Conocarpus) 
are absent from Dominica, a fact easily accounted for by the lack of suffi- 
cient lowland sites on an island where the coastline is very precipitous." 
Diligent searching by Arlington James and other members of the Division 
of Forestry and Wildlife have resulted in the verification of the occurrence 
on Dominica of three of these genera, two of which {Aviceuuia and Conocarptti) 
are reported here as new. Of the four genera listed by Hodge, only Rhiznphora 
has not yet been found on Dommica. Lagunculana was reported by Nicolson 
(1991). Only one individual o'^ Conocarpus has been found. 

St. Andrew Parish: Ballyhoo Ik-acii, Baribou Bay, Hampstt-ad, I I Dec 1993, Will & 
Jcimes 254U. (acre, CLEMS). 


Convolvnlm nodiflorus Desr. This species is widespread in dry scrub in the 
West Indies, Central and South America. It has previously been collected 
on the majority of the Lesser Antilles, including nearby Guadeloupe and 
Martiniqtie, and was to be expected on Dominica. We thank D. Nicolson 
for the identification. 

St. Joseph Parish: iMorne Raquern- (Rachctte), Au Picon Road, 1 S Dec 1 993, H/7/ 25^26 
(atrc, GH, MO, NY, US, USF). St. Peter Parish: S of Morne Espagnol between Bioche 
and Colihaut, coastal highway, 7 Jun 1996, Hill 28248 (atrc, MO, US). 


Uiiphorbui balhisii- Boiss. A species endemic to the Lesser Antilles, also known 
as Chamaesyce halbisii (Boiss.) Millsp., E. balbim was previously known only 
from Barbuda, Antigua, Guadeloupe, and La Desirade. It is a prostrate spe- 
cies of wind-swept salt-sprayed coastal cliffs in Dominica. 

St. Andrew Parish: Woodford Hill Estate, jtist E of Pte. La Sole E of an old quarry, 14 
Dec 1993, ///// &Ju»m 25464 (atrc, NY). 


Dalbergia monetarhi L.f Generally distributed from Jamaica to trojMcal South 
America, Howard (1988) reported that the species previously had been collected 
in Guadeloupe, Martinique, and St. Vincent. Velez (1 957) reported this species 
as present on Dominica, but Nicolson ( 1 99 1 ) dismissed this iis a misidentification. 
Differing from D. ecaslaphylh/n/ (L.) Taubert (which is common on Dominica 
and has unifoliolate leaves) by its pinnately compound leaves (3-5 leaflets), 
it is rather common locally along the banks of the Indian river. The identi- 
fication was confirmed by V. Rudd (SFV). 

St. John Parish: hidian River banks, Portsmouth, 5 Mar 1 99^, ///// 6 Phillippv 29056 
(atrc, BRIT, ILLS, MO, NY, US, USE, VT). 

Him. and Ja.\[F,s, Plant records for Dominica 303 


Si-da jamaicensis L. This wide-ranging species is known from Mexico, the 
Caribbean, Central America, and northern Soutli America. In the Lesser Antilles 
it has previously been collected on St. Barts, Antigua, Montserrat, Guadeloupe, 
and Les Saintes in the dry scrub forests. We thank Paul Fryxell for the iden- 

St. Joseph Parish: Morne Raquetre (Rachette), Au Piron Road, 4 Jun 1994, Hill 2580H 
(atrc, NY, TEX, USF). 


Nymphaea amazonum C.Mart. &c Zucc. This night-flowering water lily ranges 
throughout the Caribbean and from Mexico to southern Brazil. It previ- 
ously has been collected in Guadeloupe, Marie Galante, Martinique and St. 
Lucia in the Lesser Antilles. Suitable habitat for the species is very scarce 
on the island. We thank J. Wiersema for the identification. 

St. Andrew Parish: roadside 0.5 mi N ol Melville Hall Airport, Londonderry Estate, 
16 Mar 1993, Hill 24844 (atrc, CLEMS). 


Forestiera segregata (Jacq.) Krug & Urb. Another new discovery in the dry 
scrub forests of western Dominica, this species ranges from the coastal SE 
USA, S to the Lesser Antilles. It has been collected previously on Anguilla, 
Barbuda, Antigua, La Desiracle, and Marie Galante in the Lesser Antilles. 
We thank A. Goldberg and D. Nicolson (US) for the determination. 

St. Joseph Parish: Morne Raquette (Rachette), Au Piton Road, 18 Dec 1993, Hill 25530 
(atrc, GH, NY, US). 


Randia nitida (H.B.K) DC. This spiny coastal shrub is distributed from Mexico 
to South America and the Caribbean. In the Lesser Antilles it has been re- 
ported from Martinique, St. Lucia, St. Vincent, and Grenada. It is distin- 
guished from the more frequent /?. aculeata L. by its flower size (3-.3.8 cm 
vs. 2 cm long in R. aculeata) and fruit size (3.2-5 cm vs. 1-1.2 cm long in 
R. aculeata). We thank D. Nicolson (US) for the identification. 

St. Paul Parish: Rodney's Rock on the coast, Hertford Estate, 1 3 Dec 1 993, Hill 23423 


Pilocarpus race?nosus Vahl. This Caribbean endemic ranges from Cuba to the 
Lesser Antilles. It has been collected previously on Montserrat, Guadeloupe, 
and Martinique in the Lesser Antilles and was to be expected on Dominica. 
It is restricted to windswept coastal forests and is associated with the palm 
Rhyticocos on the N coast of Dominica. We thank D. Nicolson for the iden- 

304 SiDA lcS(l) 

St. Andrew Parish: clitLs at W side of Bailyboo Ik-acli, BatiboLi Bay, 1 lampsreatl, 16 
Mar 1993, Hi// 24823 (atrc, CLEMS, NY, TAHS, US, LJSF); same locality, M Dec 1993, 
///// & James 25433 (CLRMS). 


Avicennia germinans (L.) L. The black mangrove was a particularly interest- 
ing find on Dominica. While it was thought to be absent by previous au- 
thors (see comments under Cimocarpiis, above) it was located by the second 
author in recent years at several places, the most unusual one in elevated 
cliffside sedge-dominated seeps on the NE coast of the island. Other popu- 
lations are restricted to a few square meters of level land at the mouth of 
small rivers. Like Couocarpi/s, Ai'icennia is widely distributed along coasts 
from Florida and Texas, USA, to northern South America. It has been found 
on nearly all of the other Lesser Antilles. 

St. David Parish: mouth of the Ouayaperi River, Corossol Beach, 0.6 km N of La Plaine, 
12 Dec 1993, Hi// 23390 (atrc, CLEMS, BRIT, ILLS, MO, NY, US, USE, VT). St. An- 
drew Parish: Wooclhird Hill Estate, just E of Pte. La Soie E of an old quarry in sedge 
seeps, M Dec 1993, ///// OJuwes 254C^9 (atrc); motith of Eden River, Eden Estate, S of 
Crompton Point, 11 Dec 1993, Hi// & James 23473 (atrc, GH, NY, US, LJSF). 


Ctssus obovata M. Vahl. A vine endemic to the Caribbean, C. obovata is eas- 
ily distinguished from the more common C. vertiallata (L.) Nicolson & Jarvis 
by its trifoliolate rather than simple leaves. It ranges from Cuba, S to the 
northern Lesser Antilles. Collections previously have been made in the Lesser 
Antilles on St. Martin, St. Barts, and Barbuda. 

St. George Parish: head ol trail to Trafalgar Falls, Trafalgar E of hydroelectric facility, 
25 M-ay 1996, f//// 27<S7 / (atrc). 


We would like to acknowledge the continuing kind cooperation and support 
of the Ministry of Agriculture of the Commonwealth of Dominica, and es- 
pecially to the helpful staff of its Division of Forestry and Wildlife. Finan- 
cial support ior this research has been obtained primarily through various 
units of Clemson University (particularly the Archbold Tropical Research 
Center), Texas A&M University, and The LJniversity of Maine at Orono. 
The senior author was also supported by grants from the Illinois Depart- 
ment of Transportation (IDOT) and the Critical Trends Assessment Pro- 
gram (CTAP). Special thanks are extended to the late John Archbold, whose 
generosity made the research possible, to Dan Nicolson and his colleagues 
at the Smithsonian Institution lor their generous expenditure of time in 
working with several of the identifications, to our other colleagues (cited 
in the text) who assisted with identifications and collecting, and to Dr. Thomas 
Lacher, former Director of the Archbold Tropical Research Center, for his 
assistance and commitment throughout this study. 

Hill and James, Plant records for Dommica 305 


HODCE, W.H. 1954. Flora of Dominica, B.W.I. Parr 1 , Lloydia, 17:1-238. 

Howard, R.A.I 974. Flora of the Lesser Antilles, Leeward and Windward Lslands: Orchidaceae. 

Arnold Arboretum, Harvard University, Cambridge, MA. 
Howard, R.A. 1977. Floraof the Lesser Antilles, Leeward and Windward Islands: Volume 

2 Pteridophyta. Arnold Arboretum, Harvard University, Cambridge, MA. 
Howard, R.A. 1979. Floraof the Lesser Antilles, Leeward and Windward Islands: Volume 

3 Monocotyledoneae. Arnold Arboretum, Harvard University, Cambridge, MA. 
Howard, R.A. 1988. Floraof the Lesser Antilles, Leeward and Windward Islands: Volume 

4 Dicotyledoneae-Part 1. Arnold Arboretum, Harvard University, Cambridge, MA. 
Howard, R.A. 1989a. Flora of the Lesser Antilles, Leeward and Windward Islands: Vol- 
ume 5 Dicotyledoneae-Part 2. Arnold Arboretum, Harvard University, Cambridge, MA. 

Howard, R.A. 1989b. Flora of the Lesser Antilles, Leeward and Windward Islands: Vol- 
ume 6 Dicotyledoneae-Part 3. Arnold Arboretum, Harvard University, Cambridge, MA. 

NicoLSON, D.H. 1991. Flora of Dominica, Part 2: Dicotyledoneae. Smiths. Contr. Bot. 77:1- 

Velez, I. 1957. Herbaceous angiosperms of the Lesser Antilles. San German, Puerto Rico. 

Whitefoord, C. 1989. Recent plant collections from Dominica. J. Arnold Arbor. 70:l43- 



Department of Biology 

Sul Ross State University 

Alpine, TX 79832. U.S.A. 


Department of Biology 

University of Michigan 

Ann Arbor. All 48109. U.S.A. 


The widespread Old World adder's-tongue, 0/)h/oi^/oss//w poiyphyttum A. Br. is reported 
for the first time in Nortli America. Formerly confused with 0, engelmannii Prantl, we now 
recognize as 0. piilyphyll/nn collections from U.S.A. (T-xas and Arizona) and Mexico (Hidalgo, 
Oaxaca, San Luis Porosf, and Zacatecas). The earliest confirmed collection was in 1<S74. 
The two species are keyed out and illustrated, and specimens of 0. polyployllum are listetl. 
The known county distribution of the species in Texas is mapped, and the habitat and phenology 
are briefly described. 


La lengua de serpiente del Viejo Mundo, muy frecuente, Ophioglossiim polyphylli/Di A. Br. 
se cita pot primera vez en Norte America. Previamente confundida con 0. engelviannii Prantl, 
reconocemos ahora como 0. pulyphylliiin colectas de E.E.U.U. (Texas y Arizona) y Mexico 
(Hidalgo, Oaxaca, San Luis Potosi, y Zacatecas). La primera colecta confirmada fue en 1874. 
Se ofrece tma clave de identificacion y se ilustran las dos especies, y se listan los especimenes 
de 0. polyphylli/m. Se cartografia la distribucion conocida per condados en Texas, y se describen 
brevcmente el habitat y la fenologfa. 

The adder's-congues, Ophioglossum (Ophioglossaceae) are a small group 
of striking and unusual species, with the highest known chromosome numbers 
in the plant kingdom. The gametophytes are subterranean, the roots lack 
root hairs, and the leaf is made up of two parts: the sterile blade portion 
(trophophore) and the spore-bearing portion (sporophore). The trophophore 
has complex reticulate venation and is believed to be phyllodial in origin 
(Wagner 1979). The sporangia are deeply sunken in the apical portion of 
the sporophore. There are probably as many as 35 species worldwide but 
two areas, India and Africa, seem to be regions of maximum diversity. Many 
of the species occur in open arid sites and remain underground until sea- 
sonal or sporadic rains stimulate the leaves to push up through the soil and 

SiDA 18(1): 307-31.^. 1998 

308 SiDA ISd) 

grow CO full size before discharging the spores. In America norrh of Mexico 
seven terrestrial species are traditionally recognized: Ophioglossinii piisiUnni 
Raf., 0. vNli^cttuDi L., 0. ccilijoruiciini Prantl, 0. uitdicaiik L.h, 0, pdioUitiim 
Hook., 0. crotalophoroules Walter, and 0. engelvunDi'ii Prantl. However, we 
have recently discovered another species heretofore unknown from the New 
World. We originally interpreted the plants in question as a narrow form 
of 0. engelmannii (Zech & Manning 1996), but subsequent study reveals a 
number of differences from that species, and we now identify it confidently 
as 0. polyphylliim A. Br. The earliest collection was in 1874. Clausen's monograph 
(1938) does not even recognize this species anywhere in the world; he merely 
synonymized it with 0. vulgatum. Nevertheless, we have come a long way 
taxonomically from the first half of this centtiry, not only in regard to 
Ophioglossaceae, but Lycopodiaceae, Isoetaceae, Thelypteridaceae, and many 
other pteridophyte groups. Lellinger's (1985) manual of North America 
pteridophytes does not list 0, polyphyllum , nor does Flora North America 
north of Mexico (Wagner & Wagner 1993). 

This species has an enormous range: Africa, Asia (e.g., S. China, India), 
Polynesia (e.g. Hawaii), and now North America. It is probably made up of 
a number of subspecies, but these are held together by a set of distinctive 
characters, described below. There is no question that it is most closely re- 
lated to 0, eugelnunin'ii. Its main point of resemblance involves rhe venation 
pattern in which the major areoles include minor areoles, a condition com- 
monly referred to as bireticulate. In most species the major areoles of adder's- 
tongues possess only free included veinlets that themselves do not inter- 
connect. Large specimens of 0. uudicaule (the form known as 0. dlipt'icmn 
Hook. & Grev.) usually possess bireticulate veins (Wagner et al. 1984), as 
does rhe Old World 0. costcitmu A. Br. (Burrows 1996). Both Oph/ogioss//w 
viilgcitu))t and 0. ydiciilatiini show a tendency toward bireticulation, but the 
relatively few areoles are coarser and more open. The very fine bireticulation 
in 0, oigelnuinu'ii and 0. polyphyllum is cjuite distinctive. 

Ophiogloss/nii eiigeln/ciiinii is apparently a strictly New World taxon, and 
limited to North America, including Mexico, and Central America. It is 
commonly referred to as "Limestone adders-tongue" for its tendency to grow 
on soil layers above limestone. In the United States it tisually appears dur- 
ing and after rains, but it may reappear in the same places during later high 
precipitation periods. Dean P. Whittier of Vandcrbilt LIniversity has actu- 
ally forced a new "brood" of leaves to form in late summer by watering a 
site where the plants normally appear only in the spring (pers. comm.). 

A key to the two species is presented below. To see the venation of a dried 
specimen more clearly, a drop of 95% alcohol placed on the laminar surface 
will, after soaking in, clear the leaf temporarily and will not damage the 

ZncH, ET AL., A new Ophioglossum for North America 309 

Specimen. Collectors should be warned not to break off or remove the char- 
acteristic elongate brown leaf sheaths that surround the green leaf bases: 
collecting can be accomplished best by carefully digging up the specimens. 
This has the added advantage of showing the roots, and their numbers and 
relative sizes. 

Roots 25(15-30), thinner, mostly 0.6(0.3-0.9) mm thick, yellow to dark brown, 
commonly somewhat crooked (dried); persistent old leaf bases numerous, 5 
(2—9); trophophore narrowly ovate, attenuate proximally and distally, length/ 
width 4, 4.8(3.2-7.7) x 1.2(0.7-1.9) cm; sporangial clusters 1.8(0.6-2.6) 
X 0.25(0.2 -0.3) cm; apicula 1.0(0.3-1 .<S) mm O. polyphyllum 

Roots 15(6-24), thicker, 0.9-1.1(0.3-2.0) mm, often darker or blackish, usu- 
ally straight (dried); persistent old leaf bases few 2(1—4); trophophore 
broadly ovate to oblong, length/width = 3,6.0(4.5-8.5) x 2.0(1 .5-2.5); sporan- 
gial clusters 2.5(2.0-4.0) x 0.25(0.2-0.4) cm; apicula 0.6(0.0-1.3) mm. 
O. engelmannii (Fig. 1) 

The county distribution oii OphiogUnsmn polyphyllum in Texas is shown in 
Figure 2 (triangles) in comparison with 0. engelmanni {dots). The map shows 
a dramatic difference in range between the two species. Most of the collec- 
tions of 0. polyphyllum were made along U.S. Highway 90 or on roads nearby: 
in Brewster County near Alpine and Marathon; in Jeff Davis County near 
Valentine; and in Presidio County near Marfa. On the basis of our experi- 
ence, we believe that an effort ought to be made to explore much further 
afield in other Texas (as well as New Mexico and Arizona) counties, e.g., 
Hudspeth, Culberson, Reeves, Ward, Pecos, and Terrell in Texas, and also 
areas to the south in Mexico (e.g., Coahuila, Chihuahua, and Sonora states 
and southwardly adjacent states). 

There are many reasons why 0. polyphyllum was not heretofore collected, 
among the most important being its occurrence among grasses, the shape 
of the leaf resembling some monocots, and especially the very short period 
of appearance above ground, which after strong rains probably rarely ex- 
ceeds two weeks, the latter week presenting only dying straw colored and 
withered blades. 

Plants are found in shallow ditches and troughs along the roadsides (Fig. 
3). The adjacent landscape is open rangeland. While no plant species occur 
consistently with 0. polyphyllum, the two most prevalent are the grasses Buchlbe 
dactyloides and Bouteloua curtipendula. Other, less consistent associates are Bouteloua 
gracilis, Chloris verticillata, Hilaria mutica, Panicum obtusatum, and Bothriochloa 
sp. Among the forbs are Asclepias oenotheroides, Berlandiera lyrata, Cirsium 
texanum, Cooperia drummondii, CrotonVott'3Jn,Engelmanniapinnatifida, Ratibida 
columnaris, and Thymophylla sp. The vegetation is typical Chihuahuan Desert 
grassland, and includes such cactus genera as Coryphantha, Echinocereus , and 
Opuntia. The substrate involves various clays that are usually deep and allu- 

^^ 1 

SiDA 18(1) 

F1C7. 1. A semi-cliagranimatic comixinson ot (). jyil)ph)H/iiii (It-Fr) and 0, c^/^t^'t/wi/ww// (right). 
Scale bar = 2 cm. 

vial. Plants within ackler's-tongue colonies vary from several to hundreds. 
Vegetative reproduction is accomplished by horizontal roots, the prolifera- 
tions producing plantlets as far as ten centimeters from the genet. 

Our records outside of Texas are very few, and all were previously identi- 
fied as 0. engelmannii. In Arizona it was found in "mesas around the Mus- 
tang Mts." The habitats in Mexico (what little is recorded on the labels) 
appear to be more or less similar to those m Texas. The Rollins and Tryon 
collection listed below was found in "clay soil over limestone, in shade of 
small arborescent Opiiiitut in Larrea and Prosopis desert," and the Reeder et 

Zech, et al., a new Ophioglossum for North America 


Fic. 2. Presently confirmed cotinty records in Texas for 0. po!yphyll//ii/ (:riant,des) and O. 
vugelmannii (dots). 

al. collection in "thorn savannah in the open among E/ahloe dacty bides and 
Bouteloua stolonifera." The Pringle and Purpus Mexican collections lack habitat 

Collection citations: U.S.A. ARIZONA: MustanL; Mountams, 13 Sep, C. G. Pringle in 
1884, (GRAY). TEXAS. (Collections made along Hwy 90 and deposited in SRSC, tmless 
otherwise indicated. Brewster Co.: 5.5 mi E ofMarathon N of Hwy 90, 7 Jun 1995, Manning 
896; 5.9 mi W ofMarathon S of Hwy 90, 20 Jun 1995, Manning 897; 6.5 mi E of Alpine 
Sof Hwy 90, H Jul 1993. Manning 914; 12 mi W of Marathon S of Hwy 90, 5 Jul 1995, 
Mann/ng s.n.; 6.5 mi W ofMarathon N of Hwy 90, 24 Jun 1996, Mann/ng 959; 5.7 mi E 
ofMarathon, 1 1 Jun 1995. A.M. Powell & S.A. Powell 6085; 6.0 mi E ofMarathon N of 
Hwy 90, 1.^ Jun I 997, Manning 1013; 17 Jun 1997, Manning & Zech 1017 A (xVIICH); 24 
Jun 1997, Manning & Zech 10I7B (MICH); 6.2 mi E ofMarathon S of Hwy 90, 14 Jul 
1997, Hansen et al. 40; 6.3 mi S of Brewscer/Pecos Co. line, 30 Jun 1997, Manning 1034. 
Jeff Davis Co.: 7.8 mi W of Valenrine E of Hwy 90, 1 I Jul 1995, Manning 915; 25 Jun 
1997, Mann/ng & Zech 7 020 (MICH); 0.2 mi SE of 505/166 jct. on loop 166, 14 Aug 1971, 
Ke/l 1699 (GRAY); 25 Jun 1997, Manning & Zech 1031 (MICH). Presidio Co.: 0.1 mi E 
of Marfa, 26 Jul 1995, Manning 925. 


SiiiA 18(1) 

, ■-4.."BT-1|i*;'^^, 


.0. ■'^Rk"*'"^^ 



Fici. 3. Habitat of 0. polylihylli/in in sliallow roadside ditch. TX: )eff Davis C^o., alon^ loop 

MEXICO. Hidalgo: Hill, near El Ciallo, 7{)0() ft, 17 Aug 1 902 C.G. Pniii^li(GU). Oaxaca: 
Los Maranjos, vicinity of'San Lins Ttikiclanapa, Ptiebla, near Oaxaca, May 1908, C.A. Purpm 
3152 (GH). San Luis Potosi: 15 mi NE of San Luis Potosf, 6500 ft, 4 Sep 1961, _/./<?. 
Reeckr. C.G. ReeJer. & T.R. Sodentrom 3.^00 (GH). Zacatecas: 32 mi N of Fresnillo, 25 Nov 
1958, 7000 ft, R.C. RoUim ,niJ R.M. Tiyw (GH). 

There is no easily projecred certainty about when the plants will appear. 
Continuous clrout^ht will prevent the leaves from expandin^^ and extending 
above the soil. However, the best months in general arejune and July, whenever 
several rains occur. Apparently, chance rainy periods at any other time, however, 
will bring forth an "out-of-season" growth, and we now have records from 
May, August, September, October, November, and December. Our impression 
is that 0. polyphyllnni. "lies in wait" for rain, emerges through tightly com- 
pacted soil that has been softened by the water, and then has a fairly short 
time span to reproduce and disperse spores. However, based upon green- 
house observations, even when water availability is not a factor, trophophores 
will tend to die back following spore production and dispersal. 

It should be noted that 0, polyphyllimi is ecologically a "loner," in the 
sense that it does not occur microsympatrically along with other species of 
Ophioglos.u/iv (or Botrychtum). Other Ophioglossaceae are noted for forming 
"genus communities," members of the same or closely related genera that 

ZncH, ET AL., A new Ophioglossum for North America 313 

co-occur syntopically (Wagner & Wagner 1 983). To illustrate the usual situation, 
in the United States in the region from the Carolinas to Arkansas and southward, 
we find 0, nudkaule. 0. crotalophoroides , and 0. petiolatum (plus the ophioglossaceous 
Botrychium bitematum and B. lunanoides) together in the same habitat, sometimes 
side by side and intermixed with each other. But this is evidently not the 
case with 0. poly phy Hum according to Burrows (1996) who writes that in 
Africa O.polyphyllum is "The one species oi Ophioglossum that is rarely found 
as a component of genus communities." 


This study could not have been completed without the help of several 
herbaria, including Gray Herbarium, Harvard (GH), Lundell Herbarium 
(LL), University of Michigan (MICH), University of California Berkeley (UC), 
New Mexico State (NMS), Sul Ross State (SRSC), Southwest Texas State 
(SWT), Texas Tech (TTC), University of Texas at El Paso (UTEP), Botani- 
cal Research Institute of Texas (BRIT), Texas A&M University (TAMU), 
and the help of Christopher Ruggia who kindly provided the distribution 
map. Alan R. Smith and R.D. Thomas made useful suggestions for the 


Burrows, J. E. 1996. The genus Ophioglossinii L. in southern central Africa. In: J.M. Camus, 

M. GiBBY, and R.J.Johns, eds. Pteridological Perspectives. Royal Bot Gard. Kew. Pp..329-336. 
Clausen, R.T. 1938. A monograph of the Ophioglossaceae. Mem. Torrey Bot. Club. 19: 1-177. 
Clute, W. 1901 . Our ferns m their haunts. F. Stokes Co. NY. 
Lellinger, D.B. 1985. A field manual of the ferns & fern allies of the United States & Canada. 

Smithsonian Institution Press, Washington, DC. 
Wagner, W.H. 1979- Reticulate veins in the systematics of modern ferns. Taxon 28:87-95. 
Wagner, W.H., CM. Allen, and G.P. Landry. 1 984. Ophiogloisnni ellipticum Hook. & Grev. 

in Louisiana and the taxonomy of 0. nudicauk L. f. Castanea 49:99—1 10. 
Wagner, W.H. and F.S. Wagner. 1983. Genus communities as a systematic tool in the 

study ot New World Botrychium (Opiuoglossaceae). Taxon 32:5 1—63. 
Wac;ner, W.H. and F.S. Wac;ner. 1993- Ophioglossaceae C. Agardh. Adder's-tongue Family. 

In: Flora of North America Editorial C^ommittee, eds. 1993 <■. Flora of North America 

north of Mexico, 3+ vols. New York anti Oxtord. Vol. 2, jip. 85—106. 
Zech, J.C. and PR. Manning. 1996. Opbioglossi/m engelmannii in Trans-Pecos, Texas. Amer. 

Fern J. 86:102. 



Houston Comvuinity College 

1300 Holman. Houston. TX 77007, U.S.A. 


Spring Branch Science Center Herbarium 

8856 Westvieiv Drive 

Houston, TX 77033. U.S.A. 


Trinity River National Wildlife Refuge 

R 0. Box 10013 

Liberty. TX 77373. U.S.A. 


The following 15 taxaaredocLimentctI new to Texas: Alterriantlxra ses.ulis, Cyperiis ercigroUis, 
Urochloci mmosa, Lindernia criistaceci, Phyllciiith/is Irdttriu/s, Stellnria parva, Alysiairp/is vcigimdis, 
Cardamim dehilis, Rhynchospora ckhU'n, Lilaeopsis chinensis, Rumex paraguayensis, Saaharum 
coarctatum, Utriadaria foltosa, Rolygoniini mehnenaninn, and Cuiciita polygononm. Significant 
new collections records are provided for five: Sarp/is c/iheiue. Polygonum cespitos/ini, Ployllanthi/s 
iiiyiiri, Ludwtgia m'uTuairpci, ixnA Aruionnhi lacintris. Data are presented to c]i.iestion the presence 
oi Eleodoaris compressa in Texas. Taxa are listed alphabetically by himily, genus, and species 
with annotations and citation of voucher specimens. 


Los sigLiientes 1 5 taxa se citan como nnevos para Texas: Alternaiitlxra sessilis, Cyperus eragrostis, 
LJrocloloci ramosa, Lindernia irusUicen, Pl)yllcintl}//sjraternus, Stdlarici parva., Alysicarpus vciginalh, 
Cardamine de!)i!is, Rl?ynchospora dehilis, Lilaeopsis chiriensis, Rumex paraguayensis, Saaharum 
coarctat//iii, Lltric/daria foliosa, Polygoni/m meisiierniuum, y Cuscuta polygonorum. Se ofrecen nuevas 
citas importantes para cinco: Scirpus cuk'use, Polygonunj cespitiisum, Phyllanthus niruri, Ludivigia 
microcarpa, y Armoracia lacustns. Se presentan datos |-)ara ctiestionar la presencia de Eleocharis 
compressa en Texas. Los taxa estan listados altabeticamente por familia, genero, y especie 
con anotaciones y citas de los pliegos tesrigo. 


Floristic work in the Houston area has resulted m a mimeographed plant 
checklist for: Brazos Bend State Park, Fort Bend County by Larry E. Brown, 
Frank Gregg, and Kay Lewis; Davis Hill State Natural Area, Liberty County 
by Larry E. Brown, Charles D. Peterson, and Joe Liggio; and the entire Houston 
area (Harris and adjacent counties) by Larry E. Brown. In addition, the present 
authors have just completed a plant checklist for the Trinity River National 

SiDA 18(1): 315-324. 1998 

316 SiDA 18(1) 

Wildlife Refuge, Liberty County. This work coupled with plants sent to 
the SBSC^ herbarium for identification has revealed a number of plants new 
to Texas and significant new records for others. Some of these records were 
provided to Stanley D. Jones prior to the publication of his book, Vascular 
Plants oj Texas (Jones et al. 1997). Here we give data to support inclusion of 
these plants in this new publication. 

Unless otherwise indicated, all collections cited in this paper are in the 
Spring Branch Science Center Herbarium (SBSC), Houston, Texas. Dupli- 
cates of some collections are at TEX or TAES. Herbarium acronyms follow 
Holmgren etal.( 1990). 


Alternanthera sessilis (L.) DC. — A few plants of this pantropical and federal 
noxious weed were found growing in the bottomland of the Trinity River 
hi the Davis Hill State Natural Area; 4 Jul 199-1, Brown 18040. Clewell 
(1985) reported it for Escambia County in the Florida Panhandle and Tho- 
mas and Allen (1996) mapped it for ten, mostly central, Louisiana parishes. 
Nelson et al.(1997) reported it new for South Carolina and cited published 
reports for its occurrence in Alabama and Mississippi. The key in Mears 
(1977) can distinguish the Texas species oi Alternanthera, including A. sessilis. 


Lilaeopsis chtmnsis (L.) Kuntze — Affolter (1 985) mapped this species along 
the Atlantic and Gulf Coast from Nova Scotia south to Florida and west- 
ward along the Gulf Coast to Louisiana (one collection). Thomas and Allen 
(1996) mapped it for eight coastal parishes west to Vermillion Parish. The 
only Texas collection is from Galveston Bay below the Abshire House in 
the Abshire Wildlife Management Area at Smith Point, Chambers County, 
7 May 1993, Brown 16926. These plants were submerged at high tide and 
exposed during low tide. 


Arn/oracia lacustris (A. Gray) Al-Shehbaz & Bates (lake cress) — Correll and 
Johnston ( 1 970) did not report this white flowered aquatic crucifer for Texas. 
The first record appears to be in Al-Shehbaz and Bates (1987) where it is 
mapped in Tyler County. We found it to be infrequent in standing water in 
the Davis Hill State Natural Area, Liberty County (3 1 May 1996, Brown & 
Liggio 19172) and in the bottomland of the Trinity River National Wild- 
life Refuge, Liberty County (26 Apr 1 997, Brown & Marcus 20262). These 
plants produced numerous flowers but had poor fruit development. Mo- 
lecular studies based upon DNA (Les 1 994) indicate that lake cress should 
be placed into a monotypic genus Neoheckia as N. aqnatica (Eaton) Greene 
and that the cause for a poor seed set is suggested by the discovery of a trip- 
loid chromosome number of 2)i = 2H (Les et al. 1995). 

Brown and Marcus, Notes on the flora of Texas 317 

Cardamine deUlis D. Don — Plants tentatively identified as this non-native 
species have been collected in and around nurseries, greenhouses, and yards in 
the Houston area. These are similar to the native C. pennsylvanka Willd. and 
another introduced species mostly present m the norteast U.S. and Canada, C. 
flexuosa Withering. Rollins (1993) separated C. debilis from the other two species 
by its fibrillose roots and fruits <1 mm wide. Botanists may wish to reex- 
amine their collections, especially those identified as C. pennsylvanka, to see if 
they may fit the description of C debilis. It also would be helpful to search for 
additional characters to separate these three similar species. Lipscomb (1978) cited 
the Shinners collection as a voucher for the presence of C pemtsylvanica in Texas. 
Voucher specimens: Dallas Co.: weed under lath, mostly along walks in clay, Northaven 
Gardens, north Dallas, 6 Nov 1 960, Sh'niiwn 291 54 (TEX). Dewitt Co.: in Sandies Creek, 
2 mi E of Westhoff, 5 Mar 1995, D. Musthaiek j.«. (SBSC). Galveston Co.: weed in nurs- 
ery in Kemah, 1 Apr 1975, Waller & Baiiiiil 3460 (TEX). Harris Co.: weed around the 
greenhouse in the Mercer Arboretum and Botanical Gardens, off of Aldine Westheid Rd, 
I Nov 1997, Brou'/i 21713 (SBSC); growing in pots at the Edith Moore Sanctuary off of 
Memorial Avenue in Houston, 27 Mar 1992, Bri)U7/ I y86H (SBSC); weed at edge of San 
Jacinto Bldg at HCCS Central College, 1300 Holman Street in Houston, 25 Nov 1997, 
Brown 2 1742, (SBSC). 


Stellaria parva Pedersen — Landry et ai. ( 1 988) reported this chickweed new 
to North America from Acadia and Jeff Davis Parishes in Louisiana. Landry 
et al.(I989) provided a more complete discussion of its presence in Louisi- 
ana and cited an additional collection from St. Landry Parish. Thomas and 
Allen (1996) added Lafayette Parish to the Louisiana distribution. We dis- 
covered this species on a few disturbed sandy sites in the Trinity River National 
Wildlife Refuge, Liberty County, 1 1 Mar 1997, Brown & Marcus 20104; 6 
Apr 1997, Brown & Marcus 20216. Since the original description is from 
Argentina in I96I and the first Louisiana collection is in 1966, Landry et 
al. (1989) suggest it may have arrived in Louisiana and Argentina from an 
unknown native source. However, they failed to note that Pedersen (1961) 
in the protologue reported collections from Argentina and Paraguay in 1869 
and 1872 and up to 1957. 


Cuscuta polygonorum Engelmann — Correll and Johnston (1970), Hatch et 
al.d 990), Johnston (1990), and Jones et al. (1997) all include this dodder 
for Texas, apparently from Yuncker (in Lundell 1 943), who included it fide 
Small. Even though Small (1933) gave its range from Maryland to Texas, 
the first Louisiana collection of it was in 1979 with a second collection in 
1 98 1 (Gandhi & Thomas 1983; Gandhi et al. 1987). The first Texas collec- 
tion appears to be from Brazos County where it was growing on smartweed 
in the first bottomland along Highway 30, east of the Highway 6 bypass of 
Bryan/College Station, Sep 1996, Cheathaiu. Marshall & Jones s.n. 

31S SiDA 18(1) 


Cyperus eraf^rost/s Lam. — Tucker ( 19H7, 1994) reported char this species has 
become naturahzed in southeastern Texas but he cited no collections nor 
furnished details about its status in Texas. Tucker ( 1 9(S7) indicated it is native 
to the Pacific (x)ast ol the U.S., a wail in South Carolina, and naturalized in 
southern Europe. It is similar to C. ochvaceiis Vahl and may be under this 
name in otlier Texas herbaria. The key in Denton ( 1 978) can separate these 
two taxa and other similar species. The larger number of collections from 
LaPorte near the Barbours Cut Terminal of the Port of Houston suggest it 
may have arrived in the Houston area by ship. 

Voucher specimens: Galveston Co.: ca. 100 m Irom GuifOf Mexico on Iiast Beach in 
Cialvesron, 1 1 Jul 1 981 , A't'j'.i7t'/--^6J'/ (TAES). Harris Co.: in roadside diich olBarrleground 
Rd, ca, 0.5 mi S ol Hwy 225,18 Jiin 198i, llruini 1201 (SBSC); in the Marshall Tract of" 
the Ciompaq Computer Corp at the intersection of I Iwy 249 and Louetta Rd, S of Tomball, 
23 May 1992, Brown 16104 (SBSC); weedy plants in the Park Harbor development along 
Barker-Cypress Rd N ot Addicks Reservoir, W oi Hotiston, May 1 99 1, collector unknown 
(SBSC); a single plant in a prairie at end of Ramsey Rd, S of Keith Rd, E of Beltway <S in 
Pasadena, 1 Jun 199-1, Brotn/ /79^2 (SBSC); small population on verge of Barbours Cait 
Rd between hwy li6 and Barbours Cut Terminal on Galveston Bay in LaPorte, 16 ]im 
199-1, Broun ISOJJ (SBSC); frequent in ditch along Hwy 1 16 on W side across from en- 
trance to Barbours C^it Rd in LaPorte, 27 Jtm I 994, Brintii IS023 (SBSC); weedy plant at 
the Texas Parks and Wildlile Department office, near business Hwy 146 in LaPorte, 24 
Jan 1 995, Brown 1 1^403 (SBSC]); in low waste areas along Southern Pacific Railroad Tracts 
just L of Llwy 146 in LaPorte, S of the Baytown Tunnel and Bridge complex, 19 |uly 1995, 
Brotn/ /<S'6,S'9/;(SBSC). 

Eleochans a))uj>}xssci Sullivan — Correll and Johnston (1970) separate E. conipressa 
and E. aciitiiquamata Buckley, by the wider compressed culms in E. compressa 
and the more narrow non-compressed culms m E. aciitnquamatci . Further, they 
reported /:. conipressa only for San Augustine County in East Texas and re- 
stricted E. cicntisquamata to the Edwards Plateau, the north cental, and southeast 
(Refugio County) portions of the state. In addition, tliey indicated (page 275) 
that they are probably conspecific whicli is suggested by the twerlapping culms 
width measurements used to separate them in their key. We were unable to 
borrow Texas and non-Texas material ol both entities from TEX for they are 
out on loan to S. Cialen Smith for the Idora of North America project. How- 
ever, in a survey ol collections at SBSC and ASTC, we found only specimens 
referable to E. aaitisquaviata Buckley in East Texas (San Augustine and Sabine 
counties). From the specimens available to us, both entities can be readily 
separated by the nearly round culms (7-9 angled) of Z:. aciitisqiianutta that are 
up to 25 cm tall and 0.5 mm wide in contrast to the flat culms of £. compressa 
that are up to 42 cm tall and 2 mm wide. 

Rhynchospoya dehilis Gale — Gale (19-11) described this taxon as a new spe- 
cies and noted its similarity to R . fascicularis (Michx.) Vahl. Both of these 

Brown and Marcus, Notes on the flora of Texas 3 1 9 

Species were growing together at the edge of Galveston Bay on a sandy bench 
below the bluff next to the Abshire House in the Abshire Wildlife Man- 
agement Area at Smith Point, Chambers County, 7 May 1993, Brown 16932; 
27 May 1996, Brown & Liggto 19138. Here they may be distinguished by 
the taller (> 35 cm), thicker culms (> 1 .5 mm wide at culm base), and flat 
leaves (> 1.0 mm wide) oHl. fasciadciris in contrast to the shorter (ca. 15 
cm) filiform culms (< 1 mm wide at culm base) and filiform leaves (< 0.5 
wide) of/?, ckbilis. 

Sarpus cubemis Poeppig & Kunth \Oxycarynm a/Z'em^ (Poeppig & Kunth) Lye} — 
The first Texas collection of this sedge was in 1958 from Eagle Nest Lake 
m Brazoria County, Hotchktss 7633 (TEX). The second Texas collection is 
from Fort Bend County where is was somewhat frequent in a floating mat 
of vegetation in Elm Lake at Brazos Bend State Park, 7 Nov 1997, Brown 
21737 . Thomas and Allen (1993) mapped it for eight central and eastern 
Louisiana parishes, none near the Texas border. 


Phyllanthiisfraternm G.L. Webster — The first Texas collection is from downtown 
Houston growing on bare ground among planted shrubs; 17 Sep \99'b. Brown 
1 87 33. Webster (1970) indicated it is native to Pakistan and India and spo- 
radically introduced into Africa and America. He cited only three U.S. col- 
lections, one from an abandoned field in Seminole County, Florida, and the 
other two from New Orleans, Louisiana (including the first U.S. collection). 
It is now mapped for nine Louisiana parishes mostly in the New Orleans area 
(Thomas & Allen 1996). It may also expand its range in the Houston area. 

Phylianthus nin/ri L. subsp. lathyroides (Kunth) G.L. Webster — Correll and 
Johnston (1970) indicated this taxon may be extinct in Texas (the only U.S. 
records for this widespread tropical American species are from Texas) for 
they reported it only from the Ottine Swamp in Gonzales County, prob- 
ably based upon the 1935 Tharp collection at TEX. The only collections 
since 1935 are the following. 

Dewitt Co.: a group of plants ca. 2 ft tall, on sand under hridt^e on Thomaston Rd, W 
of Hwy 87 along the Guadalupe River, ca. 2 mi W of Thomaston Community, 11 Oct 
1994, Miischalek j-.k.(SBSC). Fayette Co.: S bank of the Colorado River at foot of bluff, N 
of Monument Hill State Historic Sue, 1 1 Oct 1987, Can 7867 (THX). 

The only known U.S. collections are two in the nineteenth century by 
Wright, one of which is from the Colorado River bottomland, the one in 
1935 by Tharp, and the two recent ones reported here. 


AlysicarpiLS vaginalis (L.) DC. — Isely (1990) mentions this native from the 
Old World tropics as occurring in Texas. He has examined Fabaceae collec- 

320 SiDA 18(1) 

tions at TEX and probably discovered this Hardin County specimen, 4 mi 

5 of Kountze along roadside, 22 Sep 1960, McLeoci s.n. We have collected 
plants ol this legume along a pipeline right-of-way in the Trinity River 
National Wildlife Refuge, Liberty County, 13 Oct 1996, Broivn & Marcus 


Utr/c7//ar/a/o//osa L. — This large flowered, robust, floating bladderwort was 
not reported for Texas by Correll and Johnston (1970), possibly because Small 
(1933) indicated that it ranged from Florida to Louisiana. The first report 
seems to be in Taylor (1989), where it is listed for North Carolina, Georgia, 
Florida, Mississippi, Louisiana, and Texas. However, Taylor does not indi- 
cate its distribution in Texas nor provide specimen citations. The majority 
of the TEX and LL specimens were identified as the robust U. vulgaris L. 
Taylor (1 9cS9) considers the North American representatives of U . vulgaris 
a distinct species, U . iiiacrorhiza Le (^onte, and indicates its absence from 
eastern Texas and most of the southeastern LJ.S. Coastal Plain. 

Voucher specimens: Brazoria Co.: iloaring aquatic in water in the Brazoria National 
Wildlife Refuge S of Anglcton, 26 Apr 1992, Brotrn 16020 (SBSC); Muldoon Club be- 
tween Austin Bayou and Danbury, I 9 May 1 97 1 , Fleetwood 10029 (TEX). Chambers Co.: 
present in Anahuac National Wildlife, Jul 1990, Near///e s.//.{SV,SC). Fort Bend Co.: Brazos 
Bend State Park in 40 Acre Lake beside dike between 40 Acre Lake and Pilant Lake, May 
1994, Gnxii s.n.(SBSC); 29 May 1993, Greiif^ s.n. (SBSC). Hardin Co.: 28 Jun 1952, Tharp 

6 Tyson j.^/.(TLX); Ariola Rd, S of Kountze, home of Jack Lloyd, 9 May 1957, Whitehouse 
52^HI (TEX). Llenderson Co.: Code Lake, 27 May I940, Sie^ler 2000 (TEX). Jefferson 
Co.: tloatmg in canal along rd,4 mi N of Port Authur, 19 Jul 1946, Correll 13357 (LL); 
J.D. Murphree Wildlife Mangement Area, 1-10 mi SW of Port Arthur, Big Hill Bayou, 
submerged, 20 Oct I 966, We.s/ 127 (TEX). Liberty Co.: floating on water in bayou about 

7 mi E of Cleveland, Rte. 321, 12 Sep 196S, Correll 36458 (LL). 


Ludungia /uicrocarpa Michx. — Bridges and Orzell (1989) reported this spe- 
cies new to Texas from two roadside collections in Hardin County. They 
state these collections are within former extensive wetland pine savannahs 
on the Montgomery and Beaumont Formations of the Pleistocene Epoch. 
A new county collection, perhaps from a younger formation, is from Chambers 
County where it was frequent along the sandy roadside of Highway 562 ca. 
2.8 mi east of the road to Robbins Point Park, 27 May 1996, Broivn & Liggio 



The h)llowing collections at SBSC complete the documentation of this 
species in Texas. These roadside collections suggest it may be spreading westward 
via highways. 

Voucher specimens: Hardin Co.: frecjuent in a dry roadside ditch along hwy 69/287 ca. 
1 mi N of Kountze, associated with Rbyinhnspora dheri^eiis and Viiirena hreviseta, 2 Aug 1986, 
Broun I05HI\ treqLienr in a roadside ditch at etige of a sandy acid area along Hwy 770, ca. 

Brown and Marcus, Notes on the flora of Texas 321 

0.4 mi W of intersection with Hwy 1003 between Saratoga and Kountze, 16 Aug 1986, 
Brown 1()64J', on acid sandy soil with many bog plants along Hwy 421 ca. 3 mi E of inter- 
section with Hwy 326 N of Sour Lake, 9 Jun 1996, Brown 19202. 


Saccharum coarctatum (Fern.) R. Webster — Webster and Shaw (1995) did not 
consider the presence {Erianthus) or absence {Saccharum) of an upper lemma 
awn sufficient to separate these two genera. They, tlierefore, merged Erianthus 
under Saccharum and made the appropriate combinations under Saccharum. 
They cHd not cite any Texas collections oi S. coarctatum., possibly because 
they did not examine sufficient material from Texas herbaria (the only Texas 
herbarium cited is BRIT). The following collections at SBSC document its 
presence in the state. The senior author first iclentifed these plants as E. 
brevibarbis Michx. and then annotated them to S. hrevibarbis (Michx.) Pers. 
var. brevibarbis. Joseph K. Wipff, then at TAES, identified them to S. coarctatum. 
Voucher specimens: Liberty Co.: tall plants in moist ditch along Hwy 90, 5.4 mi E of 
Hwy 61 in Devers, 25 Sep 1993, Brown 17493', tall plants with brownish inflorescence 
along Hwy 90, 4.9 mi E of Hwy 61 in Devers, 7 Nov 1994, Brown 17618. Tyler Co.: large 
colony on low moist acid soil within Kirby State Forest off of Hwy 69, S of Warren, 8 Oct 
19H5, Brown 6674. 

Uroch/oa ramosa (L.) Webster {Panicum ramostim L.; Bachiaria ramosa (L.) Stapf} — 
This native of tropical Asia has been planted in Texas and other states for 
erosion control and wildlife food but has not been included in any Texas 
floristic publications. It is reported for 31 Louisiana Parishes (Allen 1992). 
An adventive Texas collection is from Chambers County where it was growing 
at the edge of the Highway 1663 bridge over Whites Bayou northeast of 
Hankamer, 25 Sep 1993, Brown 17488. 


Polygonum cespitosum Blume — Carr and Hernandez (1993) reported this Asian 
species new to Texas (first collection from Jasper County in 1992) using 
the spelling caespitosum. We are following the spelling in Kartesz (1994). 
We discovered this species on moist to dry disturbed sites in the Trinity 
River National Wildlife Refuge. Here the reddish flowers stand out in contrast 
to the whitish flowers of P. hydropiperoides Michx. and other similar species. 
Voucher specimens: Liberty Co.: 15 Sep 1996, Brown 19701; 13 Oct 1996, Broirn & 
Marcin 19732; 9 Nov 1996, Broivn 19787; 21 May 1997, Brown 20445. The following 
two collections, previously identified as P.perskarza L., are at TAES: Galveston Co.: Galveston 
Island State Park, 15 May 1997, Rosen 0316. Liberty Co.: ca. 4.5 mi S of Dayton along 
Hwy 146, 21 Oct 1987, N/xon 16050. 

Additional state collections may be masquerading under this name in 
other Texas herbaria. 

Polygonum meisnerianum Cham. & Schlecht. var. beyrichianmn (Cham. & Schlecht.) 
Meisn. — Correll and Johnston (1970) indicated this species has been re- 

322 Sida18(1) 

ported for Texas but saw no specimens. Park (19fiS) also saw no Texas col- 
lections but reported U.S. collections from Florida and South Carolina and 
from Plaquemines and St. Tammany Parishes in Louisiana. The only Texas 
collection known to us is from Orange County where it was growing in a 
slight swale under a canopy of Chinese tallow trees adjacent to a fallow rice 
field near Nederland, Knowles sji. The senior author originally identified 
this specimen as P. siigittatuvi L. and additional collections may be filed under 
this name in Texas herbaria. 

Rumex paraguayensn Parodi — This native of southern South America (natu- 
ralized in Florida and Louisiana) is now in Texas (Chambers County, present 
in the East Unit of the Anahtiac National Wildlife Refuge off of Hwy 19H5, 
SE of Anahuac, Spring 1 99 1 , Nei'ille 274). The key in Godfrey and Wooten 
(19<^ 1 ) can separate this species from R. obovatus Danser, a similar natural- 
ized South American species in northern Florida and Louisiana. Thieret (1969) 
reported both species new to North America and speculated that R. obovcit//s 
almost certainly occurs in Texas because one collection of it was made within 
150 feet of the Texas border. However, based on Ciodfrey and Wooten's key, 
our specimen is R. paragiuiyensis. 


Lifickrnui Crustacea (L.) F. Muell. — This Indo-Malaya native ranges in the 
U.S. from South Carolina, south to Florida, and along the Gulf Coastal Plain 
to Louisiana (Godfrey & Wooten 1981). Vincent (1982) mapped it for 1 5, 
mostly south central and eastern, Louisiana Parishes. He indicated it is spreading 
westward and that it may eventually appear in East Texas. We found this 
short blue flowered species on two disturbed sites in the Trinity River National 
Wildlife Refuge, Librty County, 29 Sep 1996, Brown & Marcus 19703; 13 
July 1997, Broin/ 20581. 


We wish to thank the following; Tom Wendt (TEX), James VanKley (ASTC) 
and Stephan Hatch (TAES) for permitting access to their collections, for 
the loan of herbarium material, and for sending copies of label data from 
others; Guy Nesom and an anonymous reviewer for their careful reading 
and improvement of the paper; all of the Houston area and state-wide ama- 
teur botanists who have added collections to our local herbarium (SBSC) 
and for their help in numerotis other ways; and the Texas Parks and Wild- 
life Department and local National Wildlife Refuges for permitting col- 
lections of plants from their respective lands. Work on tlie Trinity River 
National Wildlife Refuge was sup[iorted by Challenge C^ost-Share Agree- 
ment with the refuue. 

Brown and Marcus, Notes on the flora of Texas 323 


Affoltf.r, J.M. 1985. A monograph of the genus Lilaeopsis (Umbelliferae). Syst. Bot. Monogr. 

Allhn, C.A. 1 992. Grasses of Louisiana. Second cd. Cajun Prairie Habitat Preservation Society. 

Eunice, LA. 
Al-Shhiiba/, la. and V. Ba'ii'.s. 19H7. Arnioraaa lacustris (Brassicaceae), the correct name 

for the North American lake cress. J. Arnold Arbor. 68:357-359- 
Bridcf.s, E.L. and S.L. Orzell. 1989. Additions and noteworthy vascular plant 

collections from Texas and Louisiana, with historical, ecological and geographical notes. 

Phytologia 66:12-69. 
Carr, B. and D. Hernandez. 1 993. Polygonum caes[vtos/tm var. longisetmn (Polygonaceae) new 

to Texas. Sida 15:656. 
Cleweel, A.R 1985. Guide to the vascular plants of the Florida Panhandle. Univ. Presses 

of Florida, Tallahassee. 
Correll, D.S. and M.C. Johnston. 1970. Manual of the vascular plants of Texas. Texas 

Research Foundation, Rennet, TX. 
Denton, M.F. 1978. A taxonomic cteatment of the Luzulae group oiCypems. Contr. Univ. 

Michigan Herb. 11:197-271. 
Gale, S. 1944. Rhynchospora , section Eurhynchospora , in Canada, the United States and the 

West Indies. Rhodora 46:88-134, 159-197, 207-249, 255-278. 
Gandhi, K.N. and R.D. Thomas. 1983. Cnscuta polgonorum Engelm. new to Louisiana and 

comments about its antestaminai scales. Phytologia 53:181—186. 
Gandhi, K.N., R.D. Thc:»mas, and S.L. Hatch. 1987. Cuscutaceaeol Louisiana. Sida 12:361- 

Godfrey, R.K. and J.W. Wooten. 1981. Aquatic and wetland plants of southeastern United 

States, dicotyledons. Univ. of Georgia Press, Atlncns. 
Hatch, S.L., K.N. Gandhi, and L.E. Brown. 1 990. Checklist of the vascular plants of Texas. 

Agric. Exp. Sta. MP-1655. College Station, TX. 
Holmoren,P.K., N.H. Hc:)EMc;rfn, and L.C:. BARNirrT. 1990. Index Herbariorum. Eight 

ed. New York Botanical Gatden, Btonx. 
Isely, D. 1990. Vascular flora of the southeastern United States, Vol. 3(2), Leguminosae 

(Fabaceae). Univ. of North Carolina Press, Chapel Hill. 
Johnston, M.C. 1990. The vascular plants of Texas, a list up-dating the manual of the 

vascular plants of Texas. Second etl. Ptivately published by the author. Austin, TX. 
Jones, S.D., J.K. Wipef, and P.M. Montoomkry. 1997. Vascular plants of Texas. A com- 
prehensive checklist including synonymy, bibliography, and index. University of Texas 

Press, Austin. 
Kar-im-sz, J.T 1994. A synonymized checklist of the vascular flora of the United States, 

Canada, and Greenland. 2nd ed. vol.1, (Checklist. Timber Press, Portland, OR. 
Landry, G.P., W.D. Ri;ese, and CM. Allen. 1988. Stdlaria parva Pederson new to North 

America. Phytologia 64:497. 
Landry, G.R, W.D. Reese, and K. Vincip.nt. 1 989. Stellana parva Pederson (Caryophyllaceae) 

in North America. Sida 13:373-375. 
Les, D.H. 1994. Molecular systematics and taxonomy of lake cress (Neoheck/aaq/iatica, Brassicaceae), 

an imperiled aquatic mustard. Aquatic Bot. 49:149—165. 
, G.J. Anderson, and M.A. Cleland. 1995. Sterility in the North American lake 

cress Neoheckia aquatica (Brassicaceae): inferences from chromosome number. Rhodora 


324 SiDA 18(1) 

LiPSc;()MB, l^L. 1978. Additions to rhe Texas llora. Sida 7:392-394. 

LUNDCI.L, C.L. 1943. Flora of Texas, vol. 3, part 11. Sourhern Methodist Univ. Press. Dallas, 

Mears, J. A. 1 977. The nomcncJattire and type collections oi the widespread taxa oiAltenianthera 
(Amaranrhaceae). Proc. Acad. Nat. Sci. Philadelphia 129:1-21. 

Nhi^on, J.B., K.B. Kelly, Jr., and A.C;, iVIoom;. 1 997. Noteworthy collections, South Carolina. 
Castanea 62:283-288. 

Park, Chonc-Wook. 1988. Taxonomy ot Polygouiuu section Echmocciulini (Polygonaceae). 
Mem. New York Bot. Gard. 47:1-81. 

Pedersi^n, T.M. i 9<3l . New species ai Hyclrocleis, Scirpi/s, and Stellciria. Bot. Tidsskr. 57:38- 

Rollins, R.C. 1993. The Crucilcrac oi continental North America. Stanford Univ. Press, 
Stanford, C^A. 

Small, J. K. 1 933. Manual of the sontheastern flora. Privately published by the atithor. New 
York, NY. 

Taylor, P. 1989- The ^t^enus UtriaiLir'hi-:\ taxonomic monotjraph. Kew Bull. 14:1-724. 

Thieret, J.W. 1969. Riimex obovcitiis and Riinmx pdycigiuiyoisis (Poly^onaceae) in Lotiisiana: 
new to North America. Sida 3:445-446. 

Thomas, R.D. and CM. Allen. 1993. Atlas of the vascular flora of Louisiana. Vol. 1: ferns 
and fern allies, conifers, and monocotyledons. Lotusiana Department of Wildlife & Fisheries, 
Baton Rouge. 

TiiOMA.s, R.D. and CM. Allen. 1996. Atlas of the vascular flora of Louisiana. Vol. II: di- 
cotyledons, Acanthaceae-Euphorbiaceae. Louisiana Department of Wildlife <Sc Fisher- 
ies, Baton Rouge. 

Tucker, Ci.C7 1987. The genera of (^yperaceae in the southeastern United States. J. Arnold 
Arbor. 68:361-445. 

Tucker, Cj.C 1994. Revision of the Mexican species of Cyperin (Cyperaceae). Syst. Bot. 
Monogr. 43:1-213. 

Wlb.stlr, G.L. 1970. A revision of Pb)Uct>ttl)iis (Ftiphorbiaceae) in the continental United 
States. Brittonia 22:44-76. 

Web.ster, R.D. and R.B. Shaw. 1995. Taxonomy of the native North American species of 
5^/(r/wr//w (Poaceae: Andropogoneae). Sida 16:55 1—580 

Vinc;ln'1, K.A. 1982. Scrophulariaceae of Lotiisiana, M.S. Thesis. The Lhiiversity of South- 
western Lotiisiana, Lafayette. 



U.S. Fish and Wildlife Service Missouri Department of Conservation 

608 E. Cherry St. Room 200 P.O. Box 180 

Columbia, MO 63201. U.S.A. Jeffrson City. MO 63102-0180. U.S.A. 


USD A. ARS. Southern Weed Science Research Unit 

P.O. Box 530 

Stoneville. MS 58116-0530. U.S.A. 


Stover-Ehinger Herbarium. Herbarium, Department of Biology 

Botany Department, Eastern Illinois University Valdosta State University 

Charleston. IL 61920-5099. U.S.A. Valdosta. GA 51698. U.S.A. 


Field and herbarium studies have documented Cyperus f//scus as new for Missouri and 
Nevada. Localities, habitat data, lists of associated species, description, illustrations, photo 
oF habit, and discussion of weedy potential are |iresented. 


Los estudios de campo y de herbario han documentado Cyperus fusa/s como nuevo para 
Missouri y Nevada. Se aportan localidades, datos del habitat, listas de especies asociadas, 
descripcion, ikistraciones, foto del habito, y discusion de su potencial como mala hierba. 


Cyperus fuscus L. [section Fusci (Kunth) Clarke} is native to Eurasia and 
the Mediterranean Region of northern Africa, extending from Greenland 
and Iceland to China south to Spain, Iran, Egypt, Algeria, and northern 
India (Kiikenthal 1936; Tutin et al. 1980). The common name for this spe- 
cies has been listed as "galingale," "brown galingale," or "black galingale" 
(Weedon & Stephens 1969)- It apparently was first discovered in North America 
based on specimens collected by Herbert A. Young along Revere Beach in 
Essex County, Massachusetts in 1877 (Knowlton et al. 1911; McGivney 
1938). Cyperus fuscus was subsequently documented in California (McGivney 

SiDA 18(1); 325-333. 1998 

326 SioA 18(1) 

1938; Tucker 1993), Connecticut (McGivney 1938;Dowhan 1979; Tucker 
1 995), Maryland, (Kiikenthal 1 936), Nebraska (Weedon & Stephens 1969; 
Kolstad 1986;Rolfsmeier 1995), New Jersey (Britton 1886; McGivney 1938), 
Pennsylvania {Rhoads & Klein 1993), South Dakota (Weedon & Stephens 
1969), and Virt^inia (Hitchcock & Standley 1919; Kolstad 1986; Tucker 
1987). Althougli the species has also been recorded for New York (e.g., Weedon 
& Stephens 1969; Kolstad 1986), no specimens have been located and the 
only ones so identified were actually C. diandrns Torr. (Mitchell & Tucker 
1997). Although C. J//SC//S has been present on the North American Conti- 
nent since at least 1877, its spread apparently has been slow, and outside of 
brief accounts related to its increase in distribution, there has been little 
attention <dven to it in the New World literature. 

Given the great attention to new records of Cyperus in North America by 
Charles Bryson, Richard Carter, Stanley Jones, Barney Lipscomb, Gordon 
Tucker, and others in the last 20 years (e.g., Lipscomb 1978, 1980; Tyndall 
1983; Carter et al. 1987; Carr 1 988; Carter 1988; Sundell & Thomas 1988; 
Carter & Bryson 1991; Webb et al. 1991; Bryson & Carter 1992; Jones et 
al. 1 993; Bryson & Carter 1994; Tucker 1 994; Bryson et al. 1 996; Carter et 
al. 1996), it is interesting and somewhat amazing that the species has not 
been discovered at more localities throughout the United States, especially 
as Weedon and Stephens (1969) noted that it was a weed of rice fields in 
the Old World. Outside the United States, the species has been discovered 
in Canada (GiUett 1970), but it has not been found in Mexico (Tucker 1994) 
or Central America (Tucker 1983). 


The following is a description of C fiiscin based on Kiikenthal (1936), 
McGivney (1938), Fernald (1950), Kolstad (1986), and our observations. 
Cyperiii jiisciii (Figs. 1, 2): a caesj-jitose annual with few to numerous culms 
and reddish-brown fibrous roots; culms upright, inclined, or decumbent, 
2—30 (rarely 50) cm long, 1.3—3.0 mm thick, smooth, flaccid, and triangu- 
lar in cross-section; 2—4 basal leaves per culm, 1—25 cm long, 1—4 mm wide, 
flat, flaccid, and minutely scabrous toward the acute apex; 2 or 3 leaf-like 
bracts subtend each inflorescence, varying in length, 2-25 cm long, l-3(— 
5) mm wide, spreading, flat, flaccid, and minutely scabrous toward the acute 
apex; 6— 24(— 80) densely or loosely congested spikelets per spike; spikelets 
8—1 8(— 40) flowered, 3— 8(— 12) mm long, 1 — 1 .5 mm wide, linear, and com- 
pressed; rachillas 0.3 mm wide, 0.2 mm thick, dark brown, straight or slightly 
curved, and wingless; scales (sometimes called "glumes") subtending the 
flowers O.H— 1.2 mm long, 0.8—1 .5 mm wide, broader than long, 3-nerved, 
tan or light brown at the center with narrowly hyaline margins, and the 

McKenzic, et al., Cyperus fuscus in North America 


Fic;. 1. Cyperus fuscus. A. Habit (top and side views of clump and side view of two erect 
plants). B. Inflorescence. C. Spikelet. D. Scale. E. Achene (including cross-sectional view 
and side view of 3-cleft style). (Based on McKenzie 1802 with Jacobs; McKenzie 1804; McKenzie 
1807 with Jacobs and joh>ison; illustrated by Charles T. Bryson). 

328 SiDAlSd) 

surfaces primarily dark reddish-brown to purple; the tips minutely apicu- 
late; achenes 0.9-1 -1 mm long, 0.4-0.5 mm wide, gray to almost white, 
trigonous, and narrowly ovoid; styles 0.4—0.6 mm long, 3-cleh, and not 
persistent; stamens 2 per flower, 0.7-0.8 mm long; anthers 0.4-0.5 mm 
long, tan or yellowisli, and linear oblong. 


On 9 September 1997, while conducting a search for state-listed species 
ofCyper//s, Schoenoplectus, and Lipocarpha along mudflats of the Missouri River 
in Cooper Coimty, McKenzie and Jacobs discovered a population of Cyperus 
sp. unfamiliar to them. This Cyperits was semiprostrate with the culms ra- 
diating like spokes in a wheel and leaning mostly horizontal to the ground 
{Fig. 2). The most diagnostic features of the sedge were its dark purple to 
reddish-brown scales, its strongly triangular stems, its bright rusty red roots, 
its small spikelets, and its pale, trigonous achenes. Specimens were subse- 
quently determined as Cyperm fuscin. McKenzie and Jacobs returned to the 
site on 28 September 1997 and counted 1 10 plants scattered along the silty 
mudflats and shoreline of the Missouri River, associated with the following 
species: A))i))kniui coccinea Rottb., Cyper//s diandrm Torrey, C. e)jthrorhizos Muhl., 
C. odovcttus L., C sqiicirrosiis L. (C, anstatus Rottb.; C nifhxiis Muhlenb.), 
Eclipta prostrate! (L.) L., Eragrost/s hypiioides (Lam.) Brirton, Sterns, & Pogg., 
Vi))d)rist)lis vahlit (Lam.) Link, Le/zcospora niultifida (Michx.) Nutt., Lipocarpha 
Diicrantha (Vahl) G. Tucker, VolygoniiDi spp. and Rcnuniciilin schratiis L. Al- 
though the majority of plants were in full sunlight, some extended into the 
partial shade of the developing seedlings o'l Salix spp. and Popiilus ddto'uks 
Bartram ex Marshall. 

Subsequent to this discovery, McKenzie and Jacobs were asked by Rhett 
Johnson of the Missouri Department of Conservation to identify an unknown 
specimen of Cyperin that had been collected from another site along the Missouri 
River in adjacent Howard County, approximately 32 km WNW of the original 
discovery site. McKenzie and Jacobs identified the specimen as C. fiisciis 
and visited the site with Johnson on 1 1 October 1 997, where they counted 
approximately 70 plants scattered along moist, sandy and silty flats adja- 
cent to a chute that been formed during the 1 995 flood of the Missouri River. 

Because the discovery of C, jusais in two adjacent counties along the Missotiri 
River suggested that the species could be widely distributed along the river, 
Jacobs searched locations downstream of the original discover)' site and discovered 
it in Boone, C^allaway, and Cole counties. A lack of time due to the season's 
first killing frosts prevented additional searches in other counties border- 
ing the Missouri River. With the exception of Fiiuhristylis vahlii, associated 
plants at the second, third, fourth, and fifth sites were mostly identical to 
those previously mentioned for the original discovery site. 

McKenzie, et al., Cyperus fuscus in North America 


Fic; 2. Photograph of Cyperus J //sens taken from T.isbon Bottoms, Big Muddy National Fish 
and WildHfe Refuge, Howard Co., Missouri, 1 1 Oct 1 997 (photographed by Paul M. McKenzie). 

Although the initial North American discovery of C. fuscus In Massa- 
chusetts was apparently along a sandy beach, many subsequent records along 
the Atlantic seaboard were associated with "ballast sand" (Britton 1886), 
or "waste ground, ballast, and wharf areas" (Rhoads & Klein 1993). The 
Connecticut record comes from a "sandy river shore" (Tucker 1995) and habitat 
for the species in California has been reported its "damp, disturbed soil, receding 
shorelines, and puddles" (Tucker 1993). In the Midwest, the initial discov- 
eries of this sedge were made along "wet open sandy flats" of the South Platte 
River in Lincoln County, Nebraska and "wet sandy clay soil" along the banks 
of the Keya Paha River in Tripp County, South Dakota (Weedon & Stephens 
1969). The accounts by Britton (1886) and Rhoads and Klem (1993) sug- 
gest that C. fuscus was initially introduced to the New England coast acci- 
dentally via achenes that were in ship ballast water that originated in Eu- 
rope. Introductions into other areas of the country, however, are more difficult 
to assess. It is possible that waterfowl were responsible for the species' in- 
troduction into Nebraska, South Dakota, and elsewhere. Dunn and Knauer 
(1975) postulated that waterfowl were responsible for the introductions of 
Cyperus flavkomus Mich. [C. albomarginatus (C. Martius & Schrader ex Nees) 
Steudel], Fivibristylis miliacea (L.) Vahl, and Aeschynomone indica L. into the 
Mingo National Wildlife Refuge in southeastern Missouri. 

330 SiDA 18(1) 

While the source of introduction of C. fuscus into Missouri is unknown, 
the recent and apparent widespread distribution of this sedge along the Missouri 
River suggests that the species may have become established soon after the 
1993 or 1995 floods. At the original discovery site in Cooper County, plants 
were located at the spot where a large levee break had occurred in 1995. 
Rolfsmeier (1995) reported that the two Nebraska records oiC. fusais were 
from separate locations along the Platte River. Because the Platte River empties 
into the Missouri River, and the Keya Paha eventually does the same, it is 
feasible that Nebraska or South Dakota may have been the seed source for 
the Missouri populations that became established along the Missouri River 
following the 1993 and/or 1995 floods. This introduction may be due to 
soil movement and sedimentation within the Missouri River floodplain. 

Rice growers in the United States should be alerted to the potential of 
this sedge becoming a troublesome weed. Holm et al. (1979) listed C. fuscus 
as a principal weed in Portugal and as occurring in Afghanistan and Israel. 
Based on the rapid and aggressive spread of its relative Cyperus diffornm L. 
in North American rice fields (Lipscomb 1980; Tyndall 1983; Bryson et al. 
1996), and because C. fuscus is a nee weed in the Old World (Weedon & 
Stephens 1969; Mingyuan & Dehu 1970; Holm et al. 1979), this species 
should be looked for in rice producing areas of Arkansas, California, Loui- 
siana, Mississippi, Missouri, Tennessee, and Texas. 

Due to the combination of its semiprostrate habit with the culms radi- 
ating like spokes in a wheel (Fig. 2), its dark purple to reddish-brown scales, 
its strongly triangular stems, its bright rusty red roots, its small spikelets, 
and its pale (almost white), trigonous achenes, this Cyperus is unlikely to be 
confused with any other North American member of the genus. The con- 
spicuous rusty red roots and small trigonous achenes are similar to C erythrorhizos , 
but the dark purplish scales, strongly triangular stems, apiculate achenes, 
and stamen number (2 vs. 3 in C. erythrorhizos) easily distinguish it from 
that species. The purplish-tinged scales of C. fuscus are somewhat reminis- 
cent of the scales of C, diandrus, C. bipartttus Torrey, or C. flavicovius. It can 
easily be separated from C. diandrus and C bipartitus by its trigonous vs. 
lenticular achenes, and by its smaller (ca. 1.0 mm vs. 1.8 mm) scales. It 
differs from C flavicomus by its flattened vs. erect habit, by the lack of white 
margins on the scales, and by its trigonous vs. lenticular achenes. 

The authors noted that bruised fresh and herbarium material of C, fuscus 
possesses a fragrance similar to, but not as strong as, C. scpiarrosus or C. setigerus 
Torr. & Hook. Steyermark (1963) described this odor as that of slippery 
elm {Ulm//s rubra Mtihl.). Baihl (1 995) stated that "(i)n a few genera of Cyperaceae, 
a fenugreek ifCrigonella fomuju-grciecum) odour is readily detectable in fresh 
and (especially) herbarium material." 

There is apparently some disagreement in the literature regarding the 

McKenzie, et al., Cyperus fuscus in North America 331 

number of stamens of each floret of C. jusciis. In the Vlora of the Great Flams 
(Kolstad 1986), two stamens are given in the key but three in the species 
account. One (McGivney 1938), rarely two (Kiikenthal 1936), or two (Tutin 
et al. 1980) stamens are also cited elsewhere in the literature. George Yatskievych 
(JVIissouri Botanical Garden, pers. comm.) examined all voucher specimens 
from Missouri and noted that florets had only two stamens. 

Given the apparent rapid spread of this species along the Missouri River in 
Missouri and the weedy nature of many Cyperus in the Old World, it is likely 
that this species will be discovered with sufficient effort at other Midwest loca- 
tions and possibly elsewhere in North America. While studying Cyperus speci- 
mens at the Missouri Botanical Garden in 1 988, Carter discovered a misidentified 
specimen of C. fuscus from western Nevada that originally was determined 
as C. acuminatus Torr. & Hook. The "exposed mudflat" habitat at the Ne- 
vada site is apparently similar to that in the San Joaquin Valley of adjacent 
California described by Tucker (1993). Cyperus fuscus wd& not listed byCronquist 
et al. (1977) in the Intermountain Flora, nor in Kartesz's (1987) dissertation 
on the Flora of Nevada. As with California, Missouri, Nebraska, and South 
Dakota, waterfowl may have been responsible for the introduction of achenes 
of C. Juscus into Nevada. The following data provide documentation for C. 
fuscus in Missouri and Nevada with herbarium abbreviations following Holmgren 
et al. (1990), except ctb (pers. herb, of Charles T. Bryson). 

Vouciicr specimens: U.S.A. Missouri. Cooper Co.: Big Miickly National Fish and WildHfe 
Refuge-Overton Bottoms, ca. 1.9 mi NW of Overton, T49NR1 5WSect.3 lSESENWl/4, 
Rocheport 7.5' Quad., 9 Sep 1991 , McKenzte 1802 whh Bracijacohs (ctb, EIU, MO, SWSL,VSC); 
Jacobs 97-67 with Pa/fi AkKenz/e (MO); 28 Sep 1997, McKenzw 1804 icith Brad Jacobs (ctb, 
MICH, SWSL, UMO); Howard Co.: Big Muddy National Fish and Wildlife Refuge-Lisbon 
Bottoms, ca. 1.5 mi NW of Lisbon, T50NR18WSect. 18, Arrow Rock 7.5' Quad., 11 Oct 
1997, McKenzie 1807 ivlth Brad Jacobs and Rhett Johnson (ctb, MICH, MO, SWSL, UMN, 
UMO, VDB, NSC); Jacobs 97-40 with Paul McKenzie and Rhett Johnson (ctb, MO, MICH, 
SWSL, UMN, UMO). Boone Co.: McBaine, W of Columbia waste water treatment la- 
goon # 3, T47NR13WSect. 7SWSESW1/4, Hunstdale 7.5' Quad., 13 Oct 1991, Jacobs 
97-57 with Tim Smith (MO, UMN, UMO). Callaway Co.: Notth Jefferson City, along N 
side of the Missouri River, approximately 400—500 m downstream from the Jefferson City 
bridge over the Missouri River, T44N Rl IWSect. 15SWSW 1/4, Jefferson City 7.5' Quad., 
14 Oct 1991, Jacobs 97-58 (ctb, MO, \]UO), Jacobs 97-44 with Chris Dietrich (MO). Cole 
Co.: Jefferson City, along the S edge of the Missouri River, approximately 400-500 m downstream 
from the Jefferson City Btidge over the Missouri River, just W of Wears Creek, T44NR1 IWSect. 
7NENE 1/4, Jefferson City 7.5' Quad,, 15 Oct \991 , Jacobs 97-59 (MO, NEMO, UMO); 
Jacobs 97-46 (MO). Nevada. Lyon Co.: Lahontan Reservoir, mudflats at the SE end of the 
reservoir, Tl 7NR26ESect.8, 4150 ft, plants growing on exposed mudflats, 1 Sep 1981, 
Tiehni 6769 (MO). 


We greatly appreciate the assistance of George Yatskievych, Missouri 
Department of Conservation, Flora of Missouri Project, Missouri Botanical 

332 SiDA 18(1) 

Garden, for initially identifying the Missouri specimens and providing additional 
information on C.jiiKia. We thank Connie Rutherford, U.S. Fish and Wildlife 
Service, Ventura, California, for providing information on the species' oc- 
currence in California. Steve Rolfsmeier of Milford, Nebraska, graciously 
supplied data on the Nebraska records. The helpful suggestions of George 
Yatskievych and Robert Krai improved the quality of the manuscript. 


Biui'iON, N.L. 1886. A preliminary list of N.A. species ol Cyperin, wich descriptions of 
new forms. Btill. Torrey Bot. Cllub 13:205-216. 

Biu'iii., J.J. 1995. Sedge genera ot the world: relarionshi|is and a new classiticarion of the 
Cyperaceae. Austral. Sysr. Bot. 8:125-305. 

Bryson, C.T. and R. Cakti-k. I 992. Notes on Cyper/t.s and Kyllingct (Cyi^ieraceae) in Missis- 
sippi with records of six species new to the state. Sida 15:1 19—124. 

Bkyson, C.T. and R. Carti'.r. 1994. Nt)tes on Carex, Cyperus, and Kyllinga (Cyperaceae) in 
Mississippi with records of eight species previously unreported to the state. Sida 16: 17 1 — 

Bryson, C.T.,J.R. MacDoxaii), R. (latter, and S.D.Jones. 1 996. Noteworthy Carex, Cypcr/is, 
Eleochcirii, Kyllinga, and Oxja^^v/w (Cyperaceae) from Alabama, Arkansas, Georgia, Louisiana, 
Mississippi, North Carolina, Tennessee, and Texas. Sida 17:501—518. 

(Iarr, B. 1988. Cyperiis ctifloriiiis L. (Cyperaceae) new to Texas. Sida 13:25 5—256. 

Car'i'er, R. 1988. Cypiri/s hystncnu/s (Cyperaceae) new to Florida. Sida 13:1 18-119. 

Carter, R. and C.T. Bryson. 1991- A report of Cypen/s grayioiJe.s and Cyperus retrofkxus 
(Cyperaceae) new to Missouri and notes on other selected Missouri Cyperus. Sida 14:475— 

(lARTiiR, R., C.T. Bry.son, antl B.L. Lipscomb. 1987. Cyper/is //>iiflnri/s (Cyperaceae) cast of 
the Mississippi River. Sida I 2:250. 

Car] i;k, R., R.L. Mi:ars, K.C. Bcrks, and C.T. Bry.son. 1996. A report of four exotic Cyperi/s 
((A'peraceac) species new to Lloricla, U.S.A. Sida 17:275—281. 

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Botanical Research Institute of Texas 

309 Pecan Street 

Fort Worth, TX 76102. U.S.A. 


Department of Biology 

Austin College. Sherman. TX 75090, U.S.A. 

& Botanical Research Institute ol Texas, 


A specific case of the forensic use of animal-dispersed propagiiles is presented, and it is 
suggested that this type of evidence deserves wider utilization by the hvw enforcement community. 
Animal dispersed seeds and fruits are ubiquitous, otten cling tenaciously to clothes or other 
marerials worn or used by suspects, and are small and frequently go unnoticed. Further- 
more, their identification is relatively inexpensive and technically straightforward, and their 
presentation as evidence is visually and intuitively obvious, making it ideal for the court- 
room. It is also suggested that forensic botany is an excellent topic to use as a case study in 
college botany or biology classes because of its inherent interest and integrative nature. In 
order to facilitate such usage, a brief review of some aspects of forensic botany is presented 
including references to pertinent literature. 


Se presenta un caso especffico de propagulos diseminados por animales en uso forense, y 
se sugiere que este tipo de evidencia puede tenet mayor utilizacion en varios aspectos le- 
gales. Las semillas y frutos dispersados por animales estan por todas partes, a menudo se 
enganchan tenazmente a las ropas u otros materiales llcvados o usados por sospechosos, y 
por ser pequefios pasan frecuentemente inadvertidos. ademas, su identificacion es relativamente 
barata y tecnicamente senciUa, y su presentacion como prueba es obvia visual e intuitivamente, 
convirtiendose en ideal para los juicios. Se sugiere tambien que la botanica forense es un 
tema excelente para ser usado como caso pracrico en las clases de biologfa por su interes 
inherente y naturaleza integrativa. Para facilitar ese uso se hace una breve revision de algunos 
aspectos de la botanica forense incluycndo las referencias bibliograficas pertinentes. 

The use of plants in justice and legal systems is thousands of years old, 
probably beginning in such ways as trials by ordeal (Talalaj et al. 1991; 
Mabberley 1997). In these cases, suspects were forced to eat poisonous plants 
and guilt or innocence was determined by survival. Presumably, this was 
based on a psychological effect of guilt on the vomiting reflex — suppos- 
edly, innocent individuals would expel the poisonous material, while the 

SiDA 18(1): 335-346. 1998 

SlDA 1 8( 1 ) 

guilty would retain the poison and thus die; the efficacy of such a tech- 
nique is obviously questionable. The use of plant material as evidence has 
also appeared in fictional works, such as the series of books by Ellis Peters 
about the medieval Welsh monk/herbalist/sleuth, Brother Cadfael (Whiteman 
1 995) and the works by H.C. Bailey about the detective, Reginald Fortune 
(Bailey 1936, 1943). 

The modern use of plants as evidence in a cotirt case dates to the famous 
1930s Lindbergh kidnapping case in which the son of Charles and Anne 
Morrow Lmdbergh was kidnapped and murdered. Largely through evidence 
provided by botanist Arthur Koehler, Bruno Hatiptmann was convicted of 
the crime in 1935. Koehler demonstrated that the ladder used in the kid- 
napping was built in part from wood taken from the attic of Hauptmann's 
residence (Tippo & Stern 1977; Baden 1 983; Haag 1983; Lane et al. 1990; 
Graham 1997). Koehler's evidence included identification of the wood based 
on anatomical characteristics, matching of annual growth rings, and unique 
markings made on the wood by tools including a lumber yard planer and a 
hand plane. Detailed accotints of the botanical evidence including photo- 
graphs and graphics can be found in Tippo and Stern (1977), Haag (1983), 
and Graham (1997). Graham (1997) gave an extensive list of references. 
This was a landmark case, not only because it lead to the formation of fed- 
eral kidnapping laws (Bock & Norris 1997), but also because the obvious 
value of the evidence provided by Koehler set the stage for future forensic 
uses of botanical information. 

Since that time, there have been numerous other examples of forensic 
botany (and other biological disciplines such as entomology — e.g., Rozen 
and Eickwort 1997). The use of plant fragments, pollen grains or fungal 
spores, plant trichomes (hairs), anatomical evidence from indigestible cell 
wall material from the stomach contents of crime victims, molectilar evi- 
dence utilizing DNA, and ecological evidence useful in locating hidden graves, 
crime sites, or dating when a crime occurred are just a few examples (Bock 
etal. 1988; Lane etal. 1990; IMestel 1993; Yoon 1993;Blaney 1995; Bates 
etal. 1997; Bock & Norris 1997; Graham 1997; Lewis 1997;Linden 1997). 
The following specific cases show some of the diversity in the field of bo- 
tanical forensics. Lane et al. (199(^) discussed a rape case in which leaves 
and bark fragments were found in the pants cuffs of a suspect. The material 
had gotten into his cuffs while he was climbing a tree to irain access to a 
window of the victim's house. His claim, that the victim had let him in 
through a door, did not match the botanical evidence. Another example 
used by Lane et al. (1990) involved a child abuse case. The parents claimed 
that the child had been fed fruit cocktail just prior to dying. However, their 
story was contradicted when his stomach contents showed no evidence of 
the anatomically characteristic materials expected from such a meal (e.g., 

Lipscomb and DicKis, Forensic botany 337 

stone cells from pears or needle-shaped crystals from pineapples). In the 
case of a 1989 plane crash near Ruidoso, New Mexico, it was alleged that 
an engine design flaw had allowed particulate matter (pollen) to build up 
in the engine and cause the crash. However, it was shown that smce the 
pollen was in fresh condition and had normal cytoplasm and cell walls as 
seen by electron microscopy, it could not have been exposed to the high 
temperatures present during engine operation or in the post-crash fire that 
distorted even aluminum. Further, the pollen was that of insect-pollinated 
plants found near the storage site of the plane wreckage; such pollen grains 
are unlikely to be found in any quantity in the atmosphere. Based on the 
evidence from forensic palynology, it was concluded that the pollen had gotten 
into the wreckage post-crash during several months of storage and there- 
fore had nothing to do with the accident (Blaney 1995; Brunk 1 997; Gra- 
ham 1997; Lewis 1997). A final example is the use of molecular evidence 
linking a murder suspect to a palo verde tree {Parkinsoma aadeata L., Fabaceae) 
at an Arizona crime site where he allegedly dumped the body of a victim. 
Plant geneticist Tim Helentjaris of the University of Arizona demonstrated 
that two seed pods found in the suspect's truck came from a specific palo 
verde tree scraped by the suspect's truck at the crime scene. This example is 
important because it was the first in which plant DNA was used in a crimi- 
nal case (JMestel 1993; Yoon 1993). Overviews of forensic botany were pro- 
vided by Lane et al. (1990) in the general science literature and by Bock 
and Norris (1997) in the forensic science literature. 

Our interest in this topic has developed over the course of a number of 
years during which as plant taxonomists we have been called upon numer- 
ous times by poison centers, hospitals, and law enforcement agencies to identify 
plants or their fragments. We agree with Bock and Norris (1997) that fo- 
rensic botany is a resource underutilized by the law enforcement commu- 
nity. Further, we believe that forensic botany can be very effectively used in 
botany or biology courses to show the importance, applicability, and inte- 
grative nature of botany. Because forensic botany cuts across all botanical 
disciplines and because a given case may require many research approaches 
and techniques, it is a discipline that can stress the integrative nature of 
botany and science as a whole. Additionally, it is an excellent topic to use in 
emphasizing problem solving and critical thinking skills. The purpose of 
the present article is thus two-fold. First, based on our successful use of such 
information in college teaching, we want to provide in an easily accessible 
botanical journal a brief review of forensic botany and appropriate refer- 
ences in order to encourage further such usage. Second, we present a spe- 
cific case of the forensic use of animal-dispersed propagules and suggest that 
this type of evidence has the potential to be more widely used in criminal 

3.^,8 SiDAlHd) 

On July 12, 1995, a sleeping two year old girl was pulled from the first 
floor window of an apartment in Fort Worth, Texas. The child was sexually 
molested, but fortunately was found alive about three hours later in a weedy 
area several hundred meters from where she was abducted. Assorted evi- 
dence was used in the case including fingerprints and DNA from semen. 
However, the easily understandable botanical evidence was an important 
factor in convincing the jury of the suspect's guilt. Because the Botanical 
Research Institute of Texas (BRIT) is a well known source of botanical in- 
formation in the local community, we were contacted by the district attorney's 
office to identify tiny plant fragments taken from the shoelaces of the sus- 
pect. In addition, we were provided with a bag of assorted plant material 
that had been collected from the crime scene where the child was left (Fig. 
1). Upon opening the evidence envelope, we immediately recognized the 
ca. 4 mm long plant fragments as single-seeded mericarps from a member 
of the Apiaceae (carrot family). Using a dissecting scope and authenticated 
specimens in the Botanical Research Institute of Texas herbarium, these were 
identified as mericarps oi'Tor/I/s arvensis (Huds.) Link, an introduced spe- 
cies commonly known as hedge parsley. The bag of material from the crime 
scene was then examined and a mature, fruit-bearing plant of hedge pars- 
ley was found. Under a dissecting scope, the small mericarps of this species 
(Fig. 2) are very distinctive. They are densely covered with bristles tipped 
with microscopic barbs that enhance their attachment to fur or clothing. 
They also have several very characteristic lines of closely appressed hairs between 
the bristles. Large photographs of mericarps from both the suspect's shoe- 
laces and the crime scene (Fig. 3) were presented in court by one of us (BLL). 
Like fingerprints, this was distmctive visual evidence, more easily under- 
stood than the scientifically sound but conceptually complex evidence pro- 
vided by molecular techniques such as DNA analysis. Because the suspect 
could have possibly picked up the mericarps from some other location, the 
botanical evidence alone would not have been sufficient for a conviction. 
However, it firmly linked the suspect with the crime site and in combina- 
tion with other evidence was successfully utilized by prosecutors Sharon 
McLauchlin and Larry Thompson and criminal investigator Dennis Timmons. 
The suspect, David Noel Saddlemire, was convicted of aggravated kidnap- 
ping with the jury taking only 55 minutes to reach their verdict (Fig. 4). 
He was subsequently sentenced to 99 years in the Texas state prison system. 

Ectozoochory, the transport of a diaspore or propagule on the outside of 
an animal, is a common mechanism of dispersal among flowering plants 
(van der Pjil 1982). While there are various types of diaspores (e.g., vegeta- 
tive bulbils, fragments of the parent plant), the most common types are 
seeds, whole fruits, or fruit segments (e.g., mericarps as in the above ex- 
ample). Various methods of attachment are known, ranging from viscous 

Lipscomb and Diggs, Forensic botany 339 



CASl NO. 141868 

TYFS r. i.SE Kidnapping/Sexual Assaulc Chi ld 
OFFKNSI NO. 95 364266 


SUSPSCT David Saddlemire 

ic.iv.d fromt 'l^(^/->0 ^^ L; p<.£r^^ 1^ ^'^. 


DAT! "^ • ^b TIM i I Z 3^ AM FM ^^ 


Received botanical evidence as listed below: 

A tape sealed brown paper bag containing: 

1 . A tape sealed bag holding plant material 
(II - Invoice #95 C03400) . 

2. A tape sealed envelope holding: 

A. A tape sealed envelope holding plant 
material collected from right shoe 
)?22 (Invoice ?/95C03410). 

B. A tape sealed envelope holding plant 
material collected from left shoe 
#23 (Invoice #95C03410) . 

C. A tape sealed envelope holding a 
subsample of plant material 

(28 - Invoice #95C03471). 

3. A tape sealed bag holding plant material 

(28 - Invoice //95C03471). < „ n 

~^^ PRINT GtLjgi^ Uion VQ^^Ctu 

ISLEPHONl NO. <?/^ %1^ '6D^V 


Fig. 1. Receipt of botanical evidence from the Fort Worth Police Department Crime 


SiDA lcS(l) 


Fic. 2. Torilis Lini'iisis. A) habit; B) llower; C) iruit showing mericarps with iiiicinace bristles; 
D) cross-SL'crion of frLiir (drawn by Linny Heagy). 

Lipscomb and Diggs, Forensic botany 


Left Shoe Right Shoe 

<—lcm~- > <— icm— > 


< — 1cm — > 

Fig. 3. Photographs of mericarps from botli the suspect's shoelaces and the crime scene 
(Photos by Larry A. Reynolds, courtesy of Tarrant County Medical Examiner). 

or mucilaginous substances that cause the propagule to stick to the dis- 
persal agent, to very sharp, barbed or recurved hooks, spines, or avv'ns (van 
der Pjil 1982). We believe that many of these examples are potentially valuable 
to law enforcement agencies. From many field trips both for research and 
with students, it has been our experience that it is rare to rettirn from the 
field without attached plant material either on the clothes, in pant cuffs, 
on socks, or embedded in shoelaces. Some of these are merely annoying, 
while others {e.g., Aristida species — threeawn grasses) are extremely irri- 
tating when penetrating socks or pants. Many of these seeds or fruits fre- 
quently get transferred to the interior of cars on carpeting or upholstery. 
One does not have to go far from the sidewalk to encounter such plant materials. 
Because many abundant weedy species are animal-dispersed, yards, aban- 
doned lots, and virtually any weedy or disturbed site will have some such 
species. Becatise these plant materials are so frequently encountered and because 
different plant species are found predominantly in specific habitats and during 
particular seasons, they are potentially valuable sources of evidence chat can 
link suspects with crime scenes both spatially and temporally. Also, many 
(e.g., tiny fruit segments of some Apiaceae or tiny fruits of some grasses) 


SlDA 18(1) 






FORT WORTH, TS 76196.0201 

Apnl21, 1997 

Mr. Barney Lipscomb 

Botanical Research Institute of Texas 

509 Pecan Street 

Fon Worth, Texas 76102 

Dear iVIr. Lipscomb: 

Thank you for your expertise and assistance in the David Saddlemire trial. The case was 
a complex one and your testimony was very helpful in explaining to the jury one of the 
circumstances surrounding the offense. 

The defendant was found guilty and sentenced to 99 years in the penitentiary. Again, 
thank you for your assistance. This case was an important one to the State of Texas and this 



Sharon McLauchlin 

Assistant Criminal Distnct Attorney 

Fig. 4. Letter Irom Tarrant County Olfice of the Criminal Di.strict Attorney confirming 
the jury conviction of David Noel Saddlemire. 

become deeply imbedded in cloth or carpeting, go virtually unnoticed, and 
often remain attached even after repeated washings or other types of clean- 
ing. Further, seeds and fruits are easily and inexpensively identified by trained 
botanists using nothing more than a lOX hand lens or inexpensive dissect- 
ing scope, taxonomic literature, appropriate illustrations, and herbarium 
specimens. While molecular forensic techniques can potentially provide 
very definitive information, they are much more expensive, recjuire sophis- 
ticated laboratory facilities, and are less intuitively obvious for courtroom 

Lipscomb and Diggs, Forensic botany 343 

Sometimes the attachment of ectozoochorous propagules can be quite 
tenacious. This means that they could be attached to a perpetrator's cloth- 
ing for a very long time, or even enter the flesh. The following are specific 
examples from mammals that illustrate the point. Sharp-pointed structures 
such as fruits or awns can penetrate the mouth or other tissues of livestock 
(and have to be extracted by pliers) or even become subcutaneous and re- 
quire surgical removal. Veterinarians (John Brakebill, Larry Edwards, Ken 
Lawrence, pers. comm.) indicate that it is not uncommon to find grassburs 
{Cenchrus species), awns (e.g., from Hordeum species — foxtail) or the pointed 
fruits of needle, spear, or threeawn grasses {Nassella or Aristidci species) in 
animals. For example, in North Central Texas, Cenchrus burs are often found 
embedded between the toes of dogs and grass fruits are known to penetrate 
buccal tissues including the tongue causing serious problems (Ken Lawrence, 
pers. comm.). Perhaps more striking is the ability of the sharp fruits of needle 
grass (apparently, Nassella leucotrkha (Trin. & Rupr.) Barkworth {formerly 
known as Stipa leucotrkha Trin. & Rupr.} — commonly called winter grass, 
Texas winter grass, spear grass, or Texas needle grass) to deeply penetrate 
flesh. These can become subcutaneous and require surgical removal; for example, 
they can enter between the toes of dogs and sometimes migrate long dis- 
tances subcutaneously causing draining tracts that will not heal until the 
fruit is removed (Larry Edwards, pers. comm.). The most extreme case we 
are aware of involved a fruit that penetrated through the skin and then the 
chest wall of a dog, eventually becoming embedded in a lung and causing 
a fatal case of pneumonia (John Brakebill, pers. comm.). Also tenacious are 
the spiny fruits oiTribidus terrestris L., puncture vine, of the Zygophyllaceae. 
These are very painful to both animal and human feet, damage even tires, 
and are occasionally faral to livestock if eaten (Correll & Johnston 1970); it 
would not be surprising to find them attached to various objects and pos- 
sibly even the tires of a suspect's vehicle. 

Locally in North Central Texas, we believe Soliva pterosperma (Juss.) Less., 
lawn burweed, (Asteraceae) collected from a soccer field near Arlington, Tarrant 
Co. (1995), was possibly spread from eastern Texas by athletic shoes; its 
fruit is easily, and painfully, attached by its persistent, spine-like style (Diggs 
et al., forthcoming). Such propagules could remain attached to a suspect's 
clothing or shoes for long periods of time. Other Asteraceae are well-known 
as being animal-dispersed with the pappus of many species being modified 
into an attachment structure. The retrorsely barbed awns oiBidens species, 
beggar's ticks, are strikingly effective. In another composite genus, Xanthium, 
cocklebur, the surface of the bur is conspicuously covered with stiff, hooked 
prickles ca. 5 mm long and the bur is also terminated by two prominent 
spines. The attachment of the hooked prickles to clothing or shoelaces is 

344 SiDAlSd) 

very effective and they can also easily penetrate human skin. The hooks are 
strikingly reminiscent of velcro. According to the VELCRO® Industries 
homepage (, in the early 1940s, a Swiss inventor, George 
de Mestral, after a walk noticed "cockleburrs" {presumably X.anthiuvi\ on 
his dog and his pants. He examined the hooked prickles under a micro- 
scope and derived the idea for the well known two-sided fastener — one side 
with stiff, cocklebur-like "hooks" and the other side with soft "loops" like 
the cloth of his pants. The word velcro comes from the French words ve- 
lours, velvet, and croche, hooked. 

While some of the examples above were presented to show the tenacity 
with which diaspores attach, commonly the seeds or fruits are small and 
merely cling to the fur, feathers, feet, beak, etc. of the dispersal agent with 
little or no adverse effect. Because they are often small and inconspicuous, 
they may be particularly valuable from the forensic standpoint. A well known 
such member of the Fabaceae (legume family) is the genus Desmodium, tickseeds. 
The fruits or loments are jointed and break apart into 1 -seeded flat seg- 
ments that are the dispersal units; they easily attach to hair or clothing. In 
North Central Texas for example, there are 12 members of this genus, a 
number of them occupying rather specific habitats (Diggs et al., forthcom- 
ing). Other well known examples are rhe numerous Apiaceae that have small 
schizocarps (a fruit that splits between carpels into one-seeded portions called 
mericarps) whose mericarps are covered with bristles or hairs and become 
readily attached to many surfaces. These are particularly well known to owners 
of long-haired dogs because large numbers of the mericarps become entangled 
in the fur — sometimes the situation is so severe that the only recourse is to 
shave the dog. The final example given here is the legume genus Medicago, 
commonly known as bur-clovers. There are numerous introduced species 
of this genus, many of which have prickly fruits. These fruits can be some- 
what larger than those mentioned previously, but are still effective at at- 
taching to dispersal agents. Numerous other examples could be given which 
have potential use forensically. Fortunately, most are easily identified by 
experienced field botanists because such researchers have encountered them 
many times on their own clothing or equipment. 

Other less obvious diaspores could also be potentially useful. For example, 
at the present time, several invasive aquatic species (e.g., Hydrilla verticillata 
(L.f.) Royle, in the Hydrocharitaceae) are spreading in North Central Texas, 
apparently by power boars transporting vegetative propagules (plant frag- 
ments). In areas of the country where there are numerous relatively new 
reservoirs and where the distributions of many aquatics, especially intro- 
duced species, are spotty, plant material of a given species could easily be 
used in linking a suspect with a given body of water. 

Lipscomb and Diggs, Forensic botany 345 


Forensic botany is a developing discipline that potentially has broader 
applicability than is seen at present. Technically simple, visually obvious, 
easily understood, and inexpensive methods such as the use of animal-dis- 
persed seeds and fruits are particularly worthy of further consideration. Because 
of the inherent interest in criminal cases, the potential for emphasizing problem 
solving and critical thinking skills, and the integrative nature of the sub- 
ject, forensic botany is a field that can be useful in botany and biology edu- 


We would like to thank our colleagues at various law enforcement agen- 
cies, poison centers, and hospitals for their cooperation and hard work. Special 
thanks to Dennis Timmons, Criminal Investigator of the Tarrant County 
Office of the Criminal District Attorney, and Larry A. Reynolds, Forensic 
Photographer, for the photographs, which are used courtesy of the Tarrant 
County Medical Examiner. Thanks also to John Brakebill, D.V.M., Larry 
Edwards, D.V.M., and Ken Lawrence, D.V.M., veterinarians in Sherman, 
Texas, John Lanzalotti, M.D. and Bill Vande Water, BSI, for information on 
fictional uses of plants as evidence, and Kenna Pirkle, an Austin College stu- 
dent, whose Plant Biology class paper on forensic botany prompted us to 
write this manuscript. Linny Heagy provided the illustration oiTorilis cirvtiisn. 


Badi:N, M.M. 1983. Plenary session: The LinclberL;h kidnapping revisited: Forensic sci- 
ences then and now. J. Forensic Sci. 8:1035—1037. 

Baii.IiY, H.C. 1936. A clue for Mr. Fortune: Reggie at his best. The Sun Dial Press, Garden 
City, NY. 

1943. The thistle down. Fllery Qi.ieen's Mystery JVlagazine 4:5-1 5. 

Batp.s, D.M., G.J. Ander.son, and R.D. Li.i;. 1997. Forensic botany: Tnchome evidence. J. 
Forensic Sci. 42:380-386. 

Blaney, C. 1995. Who ya gonna call:-' Bioscience 45:746-747. 

Bock, J. H., M. Lank, and D, Norris. 1 988. Identifying plant food cells in gastric contents 
for use in forensic investigations — a laboratory manual. U.S. Dej:iartment of Justice, National 
Institute of Justice, Washington, D.C. 

Bock, J. H. and D.O. Norris. 1997. Forensic botany: An under-utilized resource. J. Foren- 
sic Sci. 42:364-367. 

Brunk, S.K. 1997. The Ruidoso plane crash — the background and the trial verdict. J. Forensic 
Sci. 42:378-379. 

CoRRi;i.i,, D.S. and M.CJ. JoiixsroN. 1970. Manual of the vasculat plants of Texas. Texas 
Research Foundation, Rennet. 

Diggs, G.M. Jr., B.L. Lipscomb, and R.J. O'Kennon. Shinners & Mahler's illustrated plants 
of North Central Texas. To be published in late 1998 by the Botanical Research Insti- 
tute of Texas, Fort Worth. 

346 SiDA 18(1) 

Graiia.\[, a. 1997. Forensic palynoloi^y and the Ruicloso, New Mexico plane crash — the 

pollen evidence II. J. Forensic Science 12:391—393. 
Graham, S.A. 1997. Anatomy of the Lindbergh kidnapping. J. Forensic Sci. 42:368-377. 
Haa(,, L.C. 1983. The investigation of the Lindbergh kichiapping case. J. Forensic Sci. 

«: 1044-1048. 
LIoRNAN, J.J. 1983. The investigation ol the Lintlbergh kidnapping case. J. Forensic Sci. 

8: 1040-1 043 
Lani:, M.A., L.C. Andiirson, 'J'.M. Barkli;v, J.Ll. Bck;k, E.JVl. GitioRn, D.W. F^Al,I., D.O. 

NoRRis, TL. Ro.ST, and W.L. Sti:rn. 1990. Forensic botany. Bioscience 40:34-39. 
Li'Wis, W. H. 1997. Pollen composition in a crashed plane's engine. J. Forensic Sci. 12:387- 

LiNDi-M., CM. 1997. Field sampling and chemical analysis. J. Forensic Sci. 42:398-400. 
Mahimri.i:^', D.J. 1987. The plant book, a portable dictionary of the higher plants. Re|M-inted 

with corrections in 1989. Cambridge Univ. Press, Cambridge, Unitetl Kingdom. 
. 1997. The plant book, a portable dictionary ol the higher plants, 2nd ed. 

Cambridge Univ. Press, Cambridge, United Kingck)m. 
Me.stql, R. 1993. Murder trial features tree's genetic Fingerprint. New Scientist 138:6. 
Ro/KN, J.Ci. Jr. and G.C. Eickwort. 1997. The entomological evidence. J. Forensic Sci. 

TAi.AtAi, S., D. Talalaj, and J. 1 991 . The strangest plants in the world. iliU of 

Content Publishing Co., Melbourne, Australia. 
TiFi'o, C). and 'W.L. Stern. 1977. Fkmianistic botany. 'W.'W. Norton & Co., New York. 
VAN Der Pijl, L. 1982. Principles of dispersal in higher plants. Springer- Verlag, Berlin. 
WiiiTEMAN, R. 1995. The Cadfael companion: The world of Brother Cadfael. Mysterious 

Press Books, New York. 
YooN, C^.K. 1993. Botanical witness for the prosecution. Science 260:894-895. 



In May 1993, the senior author was informed by her friends, Don and Jo 
Robison, about an unusual population of wildflowers growing on their ranch. 
The population has been growing in the same field since the early 1960s 
when it first appeared with a culrivated batch of cotton seed (D. Robison, 
pers. comm.). The cotton was eventually replaced wirh sorghum alum and 
other grasses in order to graze cattle and the plant has continued to remain 
despite grazing upon by cattle during periods of severe drought. 

Thousands of these individuals were observed by the senior author growing 
in a sandy field. The majority of the plants, however, were toward the end 
of their blooming period and had already developed their fruits (long lin- 
ear capsules full of tiny black seeds). The senior author took the plant to be 
a poppy, however, one that she was unfamiliar with. 

This plant was later identified as Glaiiciiim corniculatum (L.) Rudolph by 
the junior author by comparison with herbarium collections at TEX. For- 
tunately, there was already a collection (0 'Kennon & Cheatham s. n. ) oiGlaucium 
from Texas. This specimen, however, was incorrectly identified (as G. flavum, 
a perennial species with yellow petals and glabrous capsules). 

In the years since 1993, Texas experienced a drought and few, if any, of 
the Glaucium in the Garza population had bloomed, preventing further 
observations by the senior author. In April 1997 sufficient rains brought 
the Glaucium population to bloom allowing the senior author to both ob- 
serve and photograph (Fig. 1) the population. Later, perhaps as a result of 
the rains, a third population v,{ Glaucium was brought to the attention of 
the junior author (Bill Henderson, pers. comm.). This population was col- 
lected at the Balcones Canyonlands National Refuge. An estimated 100 plants 
were observed growing in a circular area of about 30 feet in diameter. Two 
individuals were blooming and five were in fruit, but the majority of the 
plants were in a juvenile phase too young to flower. 

The genus Glaucium is native to Southern Europe, the Mediterranean, 
Hungary and South Central Russia. It has become adventive in California, 
Kansas, Montana, Nevada, New York, Oregon and Pennsylvania (Barkley 
1986; Hickman 1993; Kiger 1997). It is immediately distinguished from 
other Texas genera in the Papaveraceae by its fruits which are straight, unribbed, 
linear, capsules up to 25 cm long. Glaucium corniculatum is a wholly pubes- 
cent annual, 30-40 cm tall, caulescent, with clear yellow sap, cauline deeply 
pinnate unprickled leaves, inflorescence of a solitary flower that has two 
free sepals and four petals. The petals are a vivid scarlet color, two of them 

SiDA 18(1): 347. 1998 

348 SiDA 18(1) 

are wider than the other but all are oi equal length (3—4 cm). At the base of 
each petal is a purple pattern that extends toward the tip about 1/3 the 
length. The pattern is oval, 10 mm wide and lined with a creamy feather- 
like design (Fig. 1). 

Although collected in Texas as early as 1986 {Keeney 3[)()6) and recently 
reported in Texas (Jones et al. 1997), and North America (Kiger 1997), 
Glaucuint cin-iiiciihituDi has yet to be officially documented in the state. Pre- 
sented here is an updated key to the genera of Papaveraceae in Texas, a lisr- 
ing of known voucher specimens, and a photo of the flower (Fig. 1). 

Below is a revised key to the genera of Papaveraceae in the Manual of the 
vascular plants of Texas (Correll & Johnston 1970). 

1 . Acaulesccnt lu-rbs; petals 8 or more, not cTumpJL'cl in the bud; stii;mas over 

the valves ol the ca|rsLile Sanguinaria 

1 . Caulescent herbs or (ol siibacaiilescent) conlinecl to the Trans-Pecos; petals 

4 to 6, mostly crumpled in the bud; stigmas over the placentae 2 

2. Perianth and stamens borne on the rim of the hypanthiumlike expansion 
ol the receptacle; sepals united into a calyptra; Iruit conspiCLiou.sly ribbed. 


2. Perianth and stamens strictly hypogynotis; sepals not united into a calyp- 
tra; fruit not ribbed 3 

3. Herbage |-irickly; llowers on short |x'dicels; capSLile dehiscing from the 

apex by i to 6 valves Argemone 

3. Herbage not prickly; flowers on long pedicels 4 

4. Stigmatic disk present; capsules short and turgid, to ^ cm long, 
dehiscing by means of small openings jtist beneath the truncate 

summit Papaver 

4. Stigmatic disk absent; capsules linear tip to 25 cm long, dehisc- 
ing longitudinally from the apex Glaucium 

Glauciumcorniculatum(L.) Rudolph, FL Jen., 13. 1781 . CbduloniiiDicitniiaiLninu 

L., Sp. PI. 1:56. 1753. 

Voucher s|-iecimens. TEXAS. Garza Co.: community of Jiisticeburg, on harm Rd. 35 19 
on the way to Lake Alan 1 lenry, Don and jo Robison Ranch, SE corner of Ciarza C^o., ca. 3 
mi Irom Kent Co. line, in field on N side ofdirr road, 10 May 1993, Ktrkpatrick. s.ii. (TEX). 
Kerr Co.: W of Hunt on TX 39, dry creek bed on N side of road, 2 May 1 993, O'Keiinon & 
Checithcim s.n. (TEX). Travis Co.: Balcones Canyoniands National Refuge, 0.2 mi along 
dirt road NE from intersection with Cjiw (j'eek road, dirt road is 3.4 mi N along Cow 
Creek Road from 1431, just after large yellow house (30° 33' 50" N, 98" 07' 70" W), 3 
May 1991, J. K. Willuims et ul. 150 (TEX). San Saba Co.: along the Cherokee Creek on the 
William Clark (arm at Rend, on the Q'\^i:! of an over grazed field, I 1 May 1986, Keeney 
5906 (BRIT); SE of the William Clark house on the east side of the Cherokee Creek at 
Bend (1654 plants iiresent), 9 May 1987, Keony 6642 (BRIT). 

The following partial list of species from the Travis county population 

SiOA 18(1): 348. 1998 



Fic",. 1 . Photo o'i Glciucium coyniadatiaii (L.) J.H. RLiclolph. l-'hoto by Z. Kirkpacrick. 

emphasizes the weedy nature of the vegetation at this site: Argemone albiflora 
Hornem., Buglossoicks arvense (L.) I.M. Johnston*, Callirhoe involucrata (Torr.) 
A. Gray, Centaurea melitemis L.*, Cirniim texanum Buckl., Convolvulus equitans 
Benth., Ciicurbita foetidmima Kunth, Evax sp., Gauracalckola Raven & Gregory, 
Glandularia hipinnattfida (Nutt.) Umber, Lamium amplexicaule'L^', Lepidium 
virginicum L., Marruhium vulgare L.*, Medicago lupulina L.*, Hill., Oenothera 
trilobata Nutt., Physalis cinarescens (Dunal) A. Hitch., Plantago rhodospermum 
Dene., Rapistri/m rugosum (L.) AIL*, Ratihida colunmaris (Sims.) D. Don., Salvia 
farinacea Benth., Verbena halei SmaU. * = naturalized species. 
— Zoe Kirkpatrick Box 696. Post, TX 79336, U.S.A. andJ.K. Williams, De- 
partment of Botany, University of Texas. Austin. TX 78713. U.S.A. 


Barki,ey, T.M., ed. 19S6. Flora of the Great Plains. University Press of Kansas, Lawrence. 
CoRKiii.L, D.S. and M.C. Johnston. 1970. Manual of the vascular plants of Texas. Texas 

Research Foundation, Renner, Texas, 
Hickman, J.C, ed. 1993. The Jepson mantiah Hi^dier plants of C^alifornia. University of 

California Press, Berkeley. 
Jones, S., J. Wipff, and P.M. Montgomery, 1 997. Vascular plants of Texas: A comprehensive 

checklist including synonymy, bibliography, and index. University of Texas Press, Atistin. 
KiGER, R.W. 1997. Glciiicium. In: Flora North America Editorial Committee, eds. Flora of 

North America: Volume 3. Magnoliojihyta: Magnoliidaeand Hamamelidae. Oxford University 

Press, Oxford. Pp. 302-304. 

Sum 18(1): 349. 1998 

350 SiDA 18(1) 


In the lace summer of 1997 an unusual tree about 12 m tall was observed at 
the base of a high limestone cliff above Town Lake on the Colorado River in 
Austin. It had alternate leaves with prominent veins, and appeared to have 
strange fruiting structures high on the branches, these structures apparently 
deformed with galls. The tree was identified as a member of the Rhamnaceae 
based on the characteristic three-carpellate fruit with a remnant disk at the 
base, very similar to Coliibnnci. Comparison with the Rhamnaceae collec- 
tion at TEX revealed the tree to be Hovenia diilcis Thunberg (Eig. 1), the 
Chinese raisin tree, which is known to be cultivated in Texas (Jones et al. 
1997). Mabberley (1997) suggested Hovenia to be of close affinity to Colubrina, 
while recent work by Richardson (unpublished data) suggest that it is more 
closely-allied with Ziziph/is. What appeared to be galls are actually peduncles 
that swell naturally as the fruit develops. 

This species is native to mesic forests in China and was introduced into 
India and Japan for its edible peduncles (Hooker 1 872; Rehder 1 940; Roxburgh 
1975; Sargent 1916). In China the swollen peduncles of the cymose inflo- 
rescences have long been used medicinally "to offset the effects of over-in- 
dulgence in wine" (Sargent 1916). The peduncles are especially palatable 
after frost when they redden and the juice sweetens with pear-like flavor 
(Mabberly 1990; Reich 1991). The frtut itself is not edible. 

This tree is known for its cold-hardiness and is cultivated in USDA cli- 
mate zone 5, with minimum temperatures of -20"F (Dirr 1990; Staff of the 
L.H. Bailey Hortorium 1976), the equivalent of Milwaukee, WI or Binghamton, 
NY. It is therefore surprising to find it naturalized in Austin, TX, an area 
that is much warmer, and more importantly, significantly drier than much 
of zone 5. The locality where it grows in Austin is a cool, moist microcli- 
mate, at the base of a cliff of Edwards limestone and dolomite perhaps 40 
m high, facing directly to the northeast. This cliff reaches almost to lake 
level, except for a very short but steep talus slope of boulders and loamy 
soil. Vegetation growing at the base of this northeast-facing cliff is almost 
completely protected from mid-day and afternoon sun, and the presence of 
a body of water surely modifies the microclimate further. Eurther explora- 
tion via boat several weeks later resulted in the discovery of a larger tree 
perhaps 100 m upstream, and a smaller tree perhaps 100 m downstream, 
both at the base of the same cliff. Elowering specimens were later collected 
in May 1998 from the first tree discovered. 

It is possible that this small population off/, dulcis is descended from 
cultivated specimens that existed at the University of Texas College of Pharmacy 

SiDA 18(1): 350. 1998 



Fig. 1. Fruiting branch oi Hovenia diilch Tiiunb., showing capsules and swollen peduncles. 
Bar = 1 cm. 

Drug Garden, the only other specimens known from the Austin area. Prior 
to the mid- 1940s this garden existed about four miles to the north-east of 
the collection site, and was subsequently abandoned and never reconstructed 
(Beryl Simpson, pers. comm.). Two specimens exist at TEX from this gar- 
den, dated 1940 and 1946. It is possible that while H. dulcis was cultivated 
in this garden, birds ate the fruit, roosted on the cliffs and trees by the river, 
and defecated the seed to the forest soil far below. Until the early 1960's 
this portion of the Colorado river was still free-flowing, and large stretches 
of loamier soil, which is preferred by H. dukis (Dirr 1990; Staff of the L.H. 
Bailey Hortoritim 1976), likely existed. When Town Lake was createql in 
the early 1960's, perhaps a much larger population of this species was sub- 
merged and destroyed, leaving only a small number of individuals grow- 
ing near the top of the appropriate habitat. During further explorations, no 
seedlings were observed, and seedling success is probably quite low because 
numerous other species compete for space on this very thin ribbon of land. 
Furthermore, the majority of fruiting branches overhang the water, so most 

Sum 18(1): 351. 1998 

352 Si HA 18(1) 

fruit falls directly to the bottom of the lake. Similar habitats exist within 
the region, so it is possible that H. dulcn occurs unobserved elsewhere. 

Voucher specimens: TEXAS. Travis Co.: tree 1() it tall, growini; ar the base of a high, 
NE-hicmg limestone cliff along the edge of Town Lake, with Cani/is clr/niiwoiulii. PLitatms 
occidentalism and SniiUix Immi-nox, across the lake h-om the boat ramp at the University of 
Texas Brackenridge Field Lab, Austin, 30 Sep 1997, D. Goldman with J. Criitchjidd 1105 
(BLI, BRIT, TLX); same locality but with flowers, 29 May 1998, D. Goldman with P. Griffith 
1 200 (BH, BRIT, TEX). 

either specimens examined: TEXAS. Travis Co.: ciiicivated, Austin, 16 Sep I94(), C.C. 
Alhers s.n. (TEX); ex cultivation; Drug Ciarden, University ol Texas, 24 May 1946, C.C. 
All?en46ph060 (TEX). 

Thanks is given to John Crutchfield and Patrick Griffith for their assis- 
tance in obtaining specimens of this plant, and Hobbes Goldman, Beryl 
Simpson, Billie Turner, Justin Williams, and Tom Wendt for their assis- 
tance with the manuscript. — Dougla.sH. Golcliiuiu. Departmmt of Botany. Vmvmity 
of Texas. Austin, TX 78713-7640. U.S.A. 


DiRR, M.A. 1990. Manual of woody landscape plants: Their identilication, ornamental 
characteristic s, ctilture, propagation and uses. Fourth edition. Stipes Publishing, Cdiampaign. 

HooKiiR, J.l^. 1872. The flora of British India, part I. L. Reeve & CO., London. 

JoNHS, S.D., J.K. WiPrf, and P.M. Monk.omp.ry. 1997. Vasctilar |dants of Texas. A com- 
prehensive checklist. University ot Texas Press, Austin. 

MabbeiU-EY, D.J. 1997. The plant book. Second edition. Cambridge University Press, (Cambridge. 

Rehder, A. 1940. Manual of cultivated trees and shrubs. Macmillan Publishing Co., New 

Reich, L. 1991. Uncommon fruits worthy ol attention. Addison-Wesley, Reading. 

Roxburc^tH, W. 1824. Flora Indica. Reprinted by Oriole Editions, 1975, New York. 

Sakcen'I', C]. S. 1916. Plantae Wilsonianae, vol 2. Cambridge University Press, C^ambridge. 

Si'Ai'i' Ol- ■nii: L.l L Bailey Horiorium. 1976. tlortus third. Macmillan Publishing Co., New 

SiDA 18(1): ^52. 1998 

Notes 353 


The Copper Lily Habranthus tuhhpathm (L'Her.) Traub was first collected in 
Alabama in the northeast part of the city of Troy in Pike County on 1 5 July 
1988 during field work for a Flora of Pike County. This species was then 
collected in the community of China Grove in the northern part of Pike 
County on 17 July 1995 and in rural western Bullock County near the area 
referred to as High Ridge on 12 July 1996. A fourth site southwest of the 
city of Troy in Pike County was discovered in August of 1997 (Fig, 1). These 
are the first records of this species from Alabama, and the second record of 
this species in North America outside of Texas and Louisiana according to 
Holmes and Wells (1980) and Burkhalter (1984), 

Habranthus tuhispathus (L'Her.) Traub (Liliaceae), also known as Zephyrantlm 
texana Herbert or Habranthus texanus (Herbert) Herbert ex Steud,, is a sea- 
pose perennial from a small bulb. Each 10-20 cm scape bears a single yel- 
lowish-orange flower produced during the summer months, usually following 
a rain. The leaves are linear, 3-5 mm wide and up to 25 cm in length, and 
appear in autumn and wither by early spring. The fruit is ovoid and three 
lobed containing numerous flat, black papery seed at maturity. The genus 
Habranthus is separated from the genus Zephyranthes by Sealy (19.37) based 
upon the filaments oi Habranthus being of four lengths and the anthers af- 
fixed at the middle, while in Zephyranthes the filaments are of alternate lengths 
and the anthers affixed below the middle. 

In the city of Troy, Copper Lily is found growing in lawns, along side 
walks, in a city softball field, a grass parking lot, and a cemetery. All of the 
sites are to some degree disturbed and other vegetation consists primarily 
of mixed grasses (predominately Cynodon dactylon (L.) Pers., Paspalum notatum 
Fluegge, and Eremochloa ophiuroides (Munro) Hack.) and other bulb species 
Qpheion uniflorum (Lindley) Raf , Allium bivalve (L.) Kuntze, and Allium canadense 
L,), Copper Lily is so abundant there that in some lawns it forms a com- 
plete ground cover when in foliage. The soils are sandy and well drained. 
Flowering of Copper Lily is most common in July, August, and September, 
when periods of drought are broken by summer thunder showers. The foli- 
age appears in late autumn, usually in October, and remains green throughout 
the winter. The total area occupied by Copper Lily in Troy is about three to 
five acres, and extends in a unbroken population for 276 meters along Three 
Notch Street. 

In China Grove and in the High Ridge area Copper Lily is found on roadsides 
and in lawns. It is associated with many of the same species as the popula- 
tion in Troy, with Cynodon dactylon (L.) Pers. and Paspalum notatum Fluegge 

Sum 18(1): 353. 1998 


Sum 18(1) 

Fig. 1. Map ot Alabama with locations of Pike and Bullock counties. 

predominating. The soils in Cliina Grove and in High Ridge are coarse sands 
and the sites tend to be xeric. Undisturbed areas nearby support a dwarf 
oak-lichen sandhill community. The population in China Grove extends 
for 301 meters along Pike County Highway 37 and the High Ridge popu- 
lation extends for 142 meters along Bullock County Highways 7 and l4. 
Estimates of total population size are three acres for China Grove and two 
acres for High Ridge. 

The fourth population is located about eight miles southwest of Troy 
near Goshen. It is also found on a roadside and extends for 75 m along Pike 
County Highway 25. The soil is sandy and the main associate at this site is 
Paspalum notatum Fluegge. This is the smallest population, covering approximately 
one-hall acre. 

Monitoring of a planted population of copper lily for five years revealed 
that the seed generally fell within 15 cm of the parent plant and that seed- 

SiDA 18(1): 354. 1998 

Notes 355 

lings were readily produced. Seedlings generally flowered for the first time 
in their third year of growth without special care. 

Holmes and Wells (1980) reinforced the suggestion of H.H. Hume (Sealey 
1937) that the distribution of Copper Lily in North America was the result 
of human activity. They speculated that the distribution of the Copper Lily 
in North America was most likely the result of traveling Spanish mission- 
aries as all of the Texas and Louisiana populations are near the sites of Span- 
ish missions. As there is no record of a Spanish mission ever having been 
located in Troy or in Pike county, it is most likely that the species reached 
Alabama by other means. Historical records of the Troy-Pike county area 
reveal that in 1865 a man by the name of William Murphree left Troy and 
moved to Walker County, Texas. In letters to his mother, Murphree states 
that he is enclosing seeds for her to plant. In other records Mrs. Murphree 
is noted for planting many of the decorative plants in Troy at that time. 
The Murphrees lived on Three Notch Street in Troy, the site of the largest 
Copper Lily population reported above. Walker County, Texas, where Mr. 
Murphree moved is listed by Holmes and Wells as a place where Copper 
Lily was collected (1980). We therefore speculate that Copper Lily was in- 
troduced into Alabama by the Murphree family from populations in Walker 
County, Texas. As all of the sites reported above are old communities, it is 
likely that Mrs. Murphree shared the plant with friends or relatives nearby. 

Voucher specimens: ALABAMA: Bullock Co.: T13N R21-22E sect. 36-31, intersec- 
tion ot Bullock Co. Hwy 7 and 14 and un-numbercd dirt road, high ridge, roadsides and 
lawns, hill sun, deep dry sandy soil, abundant, 12 Jul 1996, A.R. Diamond 10413 (AUA). 
Pike Co.: TION R21E sect. 29, Troy, U.S. Hwy 29 (Three Notch Street), north of down 
town at Knox Field, abundant along sidewalks and road, lawns, and vacant lots, full sun, 
sand or clay soil, flowered after shower during drought, no leaves seen, 15 Jul 1988, A.R. 
Diamond 5210 (AUA); T12N R21E sect. 9, China Grove, just east of Pike Co, Hwy 7, 
roadside, full sun, dry sandy soil, common, 17 Jul 1995, A.R. Diamond 9691 (AUA); T9N 
R20E sect. 6, Pike County Hwy 25, 0.2 mi S of U.S. Hwy 29, ca. 75 m along rhe west side 
of the road, full sun, flowers yellow, 3 Oct 1997, A.R. Diamond 11131 (AUA). 

Duplicates will be distributed at a later date. — Alvin R. Diamond, Jr., 

Charles P. Chapman, and Jim Brummett, Department of Biology, Troy State Uni- 
versity, Troy, AL 36082. U.S.A. 


BuRKHALTER, J.R. 1984. Additions to the vascular flora of Florida. Castanea 49:181. 
Holmes, W.C. and C.J. Wells. 1980. The distribution oi Hahranthus tubispathus (L'Her.) 

Traub in South America and Norrh America- Texas and Louisiana. Sida 8:328-333. 
Sealy, J.R. 1937. Zephyranthes, Pyrolirion, Hahranthus and Hippeastrum. J. Royal Hort. Soc. 


SiDA 18(1): 355. 1998 

356 SiDA 18(1) 


Plantago coronopus L,, native to Europe and the Mediterranean area (Chater 
& Carrier 1976), was reported as naturalized in California (Dempster 1993). 
In the eastern United States it has been variously described as rarely adven- 
tive on ballast heaps at seaports (Gleason 1952), occurring sporadically about 
ports without persisting (Fernald 1950), and as a repeated introduction in 
ballast that does not become established (Gleason & Cronquist 1991)- It 
was not reported in standard floras of the southeastern United States (e.g., 
Small 1913; Radford et al. 1968), or in the floras of Missouri (Steyermark 
1963), New Mexico (Martin & Hutchins 1981), Great Plains (McGregor & 
Brooks 1986), Arkansas (Smith 1994), Oklahoma (Taylor & Taylor 1994), 
or Texas (Correll & Johnston 1970). It was also not included in more recent 
works on Texas plants such as Hatch et al. (1990) or Jones et al. (1997), nor 
was it listed by Rosatti (1984) in his treatment of Plantaginaceae for the 
Generic Flora of the Southeastern United States. 

As part of the collecting effort for the forthcoming Shinmrs & Mahler's 
Illustrated Flora of North Central Texas (Diggs et al., forthcoming), a collec- 
tion made in 1998 in Fort Worth (Tarrant County) is apparently the first 
documented occurrence of this species for Texas. 

Voucher specimen: TEXAS. Tarrant Co.: weedy area in landscape, 301 Cresrwood, Fort 
Worth, 24 May 1998, O'Keumm 14221 (BRIT), 

At the collection locality numerous individuals were observed as landscape 
weeds and other individuals were observed in nearby weedy areas. The mode 
of introduction is unclear, but seeds were possibly introduced with landscape 
plants. Introduction could also have occurred through spread from intentional 
cultivation. Since 1997, seeds oiPlantago coronopus have been offered by Johnny's 
Selected Seed Company for the specialty salad green market under the name 
"Minutina" or "Erba Stella" and, until recently, seeds were sold by the Orna- 
mental Edibles Seed Company as an "edible landscape plant." Other north 
central Texas populations have not been found and it is unclear whether the 
species will become more widely established. However, some individuals were 
very robust, with one having 65 inflorescences. Also, the species is self-com- 
patible (Delden et al. 1992) and is tolerant to drought, soil compaction, and 
competition (Mook et al. 1992). The seeds are highly viable and germinate 
well in soils with low moisture content (Blom 1992). 

This species, commonly known as buck-horn plantain, differs from all 
other plantains occurring in Texas in having deeply pinnately lobed leaves, 
short hairs on the corolla tubes, and 3- or l-locular capsules. Other mem- 
bers of the genus occurring in the state have leaves entire or with few teeth 
or shallow lobes, corolla tubes glabrous, and 2-locular capsules. 

SiDA 18(1): 356. 199<S 



Fig. 1. Habit of Piantago corunopus (with permission from Gleason 1952). 

Plantago coronopus (Fig. 1) can be recognized by the following descrip- 
tion (Butcher 196l;Gleason 1952; Chater& Carrier 1976; Dempster 1993; 
Rahn 1996): Pubescent annual or biennial with a taproot and leaves in a 
basal rosette; leaves closely spreading on the ground or ascending, linear to 
lanceolate in outline, 4—25 cm long, with spreading-ascending, acute, ± 
linear lobes, the lobes sometimes toothed or lobed; inflorescences sometimes 
numerous, 5—50 cm long including the leafless, densely hairy peduncle terminated 
by a narrowly cylindric, dense spike to ca. 12.5 cm long; bracts usually not 
surpassing the flowers, broadly scarious-margined at base, the keel prolonged 
into an acuminate tip; corolla tubes covered with short hairs; petals acute; 
stamens 4; capsules 3- or 4-locular; seeds 3—5 per capsule, dimorphic, one 
per locule, ca. 1.5 mm long, and one smaller than the others, ca. 0.5 mm 
long, occurring in an upper compartment in the ovary on the abaxial side; 
flowering late Apr— Sep. 

— Robert J. O'Kennon. Botanical Research Institute of Texas, 509 Pecan Street, 
Ft. Worth, TX 76102, U.S.A.,; George M. Diggs,Jr., De- 
partment of Biology, Austin College, Sherman, Texas 73090, U.S.A. & Botanical 
Research Institute of Texas,; and Ronald K. Hoggard, Department 
of Biology, University of Central Oklahoma, Edmond, OK 7 5034. 

SioA 18(1): 357. 1998 

358 Si DA 18(1) 


Bi.OM, C.W.P.M. 1992. Ecology oi Plantci^o populations: Germination and establishment. 
In: RJ.C;. KtnixT & M. Bos, eds. PLnilago: A Mukidisciplinary Sttidy. Pp. 88-97. Springer- 
VerLag, Berlin, Heidelberg, New York. 

Bu'rciiiui, R.W. 1961. A new illustrated Britisli flora. Leonard Hill Books Limited, Lon- 
don, England, U. K. 

C()KK];i.[., D.S. and M.C^. Johnston. 1970. Manual ot the vascular plants ot Texas. Texas 
Research Foundation, Rennet. 

CiiA'iRR, A.O. and D. Cartif.r. 1976. P/a//tc/go. In: T.G. Turin, V.H. Heywood, N.A. Burges, 
D.M. Moore, D.H. Valentine, S.M. Walters, and D.A. Webb, eds. Flora Europaea 4:38-44. 

Di-LDiiN, W. Van, 1 1. Van Dijk, and K. WoLif. 1 992. The genetics o^Plantnii^o species. In: 
P.J.C. KuiperandM. Bos, eds. PLnitagi): A mukidisciplinary sttidy. Pp. 295-309. Spnnger- 
Veriag, Berlin, Heidelberg, New York. 

Drmpsthr, L.T. 1993. Plantaginaceae. In: J.C.Hickman, ed. The Jepson manual, higher 
|ilants of C^alifornia. Pp. 820-82 I . Univ. of (California Press, Berkeley. 

DiGCiS, G.M. Jr., B.L. Lipscomh, and R.J. O'Kt.nnon. Shinners & Mahler's illustrated flora 
of North Central Texas. To be published in late 1998 by the Botanical Research Insti- 
tute of Texas, Fort Worth. 

Ft;RNAi.i), M.L. 1950. Gray's manual ol botany, 8th ed. Reprinteti I 987. Dioscorides Press, 
Portland, OR. 

Gi.i;ason, H.A. 1952. The new Bntton and Brown illustrated flora ol the northeastern United 
States and adjacent Canada, 3 vols. New York Botanical Garden, Bronx. 

anti Cronqulsi', A. I 99 1 . Maniial ol the vascular plants of northeastern United 

States and adjacent Canada, 2nd ed. Van Nostrand Reinhold Company, New York. 

Hatch, S.L., K.N. Gandhi, and L.E. Brown. 1 990. Checklist of the vascular plants of Texas. 
Texas Agric. Exp. Sta. Misc. Publ. 1655:1-158. 

Jt)Ni;s, S.D., J.K. WiPif, and P.M. Montoomi-rv. 1 997. Vascular plants of Texas: A compre- 
hensive checklist including synonymy, bibliography, and index. Univ. of Texas Press, Austin. 

Makiin, W.C. and C.R. Hutchins. 1981. A flora oi New Mexico, 2 vols. J. Cramer, Ger- 

McCiKHCOR, R.L. and R.E. Brooks. 1986. Plantaginaceae. In: (ireat Plains Elora Associa- 
tion. Flora of the Great Plains. Pp. 742-747. Univ. Press ol Kansas, Lawrence. 

Mook,J.H.,J. HaI'Ck.J. Vandf.r Took n, and PH. van Tif.ndi;ri.n. 1992. Ecology of /Vrf;//t/,(;« 
populations: The tlemographic srructure of iioj-iulations. In: P.J. (7 Kuipcr and M. Bos, 
eds. Plantago: A mukidisciplinary study. Pp. 69-87. Springer- Verlag, Berlin, Heidel- 
berg, New York. 

Radiord, A.E., H.E. Ahlhs, and (7R. Bi-ll. 1968. Manual of the vascular flora of the Caro- 
linas. The Ihiiv. of North Carolina Press, Chapel Hill. 

Raiin, K. 1996. A phylogenetic study of the Plantaginaceae. J. Linn. Soc, Bot. 1 20: 145- 

RosAiii, T J. 198 1. The Plantaginaceae in the soLitheastern United States. J. Arnold Arbor. 
65: 533-562. 

Small, J.K. 1913- Flora of the sotitheastern United States, l\^^ etl. Published by the au- 
thor. New York. 

Smiiii, E.B. I99I. Keys to the flora of Arkansas. Univ. of Arkansas Press, Fayetteville. 

Sii;vi;RMARK, J.A. 1963. Flora of Missouri. The Iowa State Univ. Press, Ames. 

Taylor, R.J. and C.E.S. TAMxm. 1 994. An annotated list of the ferns, fern allies, gymnosperms 
and flowering plants of Oklahoma, 3rd ed. Southeastern Oklahoma State Univ., Durant. 

SiDA 18(1): 358. 1998 

Notes 359 


Pteris vittata L., commonly known as ladder brake, Chinese brake, or Chi- 
nese ladder brake, is an Asian native well known as an escape from cultiva- 
tion in coastal areas of the southeastern United States. The species was not 
included in Small's 1903 or 1913 treatments of the southeastern flora. Small 
later (1938) treated the species as Pycnodoria vittata (L.) Small, indicating 
that it had been collected in Florida many years previously and that during 
the 1930s it was abundant at several Florida localities (e.g.. Everglades). 
He also indicated that it was established in Alabama, Louisiana, the West 
Indies, and South America. Currently, Pteris vittata is common in southern 
Florida where it can be found in almost every habitat with exposed lime- 
stone (e.g., pinelands) and on a variety of man-made calcareous substrates 
(e.g., sidewalks, buildings, old masonry) (Nauman 1993). Radford et al. 
(1968) cited a South Carolina record and noted that it was a rare escape. 
Nauman (1993) mapped the species as occurring from southern South Carolina 
south to Florida and west to eastern Louisiana; he also showed isolated oc- 
currences in southern California and the District of Columbia. 

Pteris vittata was not included in the Texas flora by Correll (1956, 1966), 
Correll and Johnston (1970), Stanford (1976), Hatch et al. (1990), or Jones 
et al. (1997), nor has it been reported from Oklahoma (Taylor & Taylor 1994). 
It is now known from Texas based on the following collection: 

Voucher collection: Texas. San Saba Co.: ca. 3 mi N of Bend, on the Edwards Plateau, 
growing from a limestone boulder with Adiantuni capillns-veneris L. at edge of Rough Creek 
(tributary of the Colorado River), 6 Nov 1987 J. W'! Stanford 5 iOS (BRIT, HPC, SPLT). 

Subsequent flooding has destroyed the site. The identification was con- 
firmed by Jim Blassingame (South Plains College, Levelland, TX). 

Of the five Pteris species in the United States (most occurrences are in 
the southeastern part of the country), four are introduced and one, P. bahamensis 
(J. Agardh) Fee, is native to south Florida (Nauman 1993). Besides P. vittata, 
only P. multifida Poir. (spider brake, Chinese brake, Huguenot fern), is known 
from Texas. This widely cultivated native of China is naturalized in east 
and southeast Texas and the Rio Grande Valley (Correll & Johnston 1970; 
Hatch et al. 1990). The deeply palmately 3-divided pinnae (at least some) 
of P. multifida are quite distinctive. 

Pteris vittata can be recognized and distinguished as follows: stems short- 
creeping, stout, densely scaly; leaves clustered, strictly 1 -pinnate (the pin- 
nae without lobes or divisions), 0.3-1 m long including petiole; pinnae 12- 
20(-30) pairs per leaf, linear-lanceolate to linear-attenuate, 2-18 cm long, 
4—9 mm wide, long-attenuate or sharply acute apically, asymmetrically cordate 
to widened or truncate basally; serrulate marginally; distal pinnae conspicuously 

SiDA 18(1): 359. 1998 

360 SiDAl8(l) 

longer than proximal pinnae (leaf blades thus oblanceolate in outline) with 
the terminal pinna typically longest; petioles and often rachises densely scaly; 
sori usually of a continuous narrow band near the margins of the abaxial 
surface of the pinnae (thus most of the abaxial blade surface exposed) (Long 
& Lakela 1971; Nauman 1993; Smith & Lemieux 1993). The species is a 
tetraploid, with 2« =116 (Nauman 1993). An illustration and a detailed 
description are available in Small (1938). A recent treatment oi Pteris, in- 
cluding a key to taxa occurring in the United States and an illustration of 
P. vittata, can be found in Nauman (1993). 

— -Jack W. Stanford. Department of Biology. Howard Payne University, Brownwood, 
TX 76801. U.S.A.. afui George M. Diggs,Jr, Department 
of Biology. Austin College, Sherman, TX 75090. U.S.A. & Botanical Research 
Institute of Texas, 509 Pecan Street, Ft. Worth. TX 76102, U.S.A., 


CoRRJiLL, D.S, 1956, Ferns and fern allies of Texas, Texas Research Foundation, Rc-nncr. 

1966, Pteridophyta, In: C.L, Lundell, ed. Flora of Texas 1:3—121. Texas 

Research Foundation, Renner. 

and M.C,, 1970, Manual of the vascular plants of Texas, Texas 

Research Foundation, Renner, 
Hatch, S,L., K,N. Gandhi, and L.E, Brown, 1 990. Checklist of the vascular plants of Texas. 

Texas Agric. Exp, Sta. Misc. Publ, 16^5:1-158, 
JoNF.s, S,D,,J,K, Wii'i'i^and P,M, Mon'k.omI'UY. 1997. Vascular plants of Texas: A comprehen- 
sive checklist including synonymy, bibliography, and index, Univ. of Texas Press, Austin, 
IxjNG, R.W, and O. Lakela. 1971, A flora of tropical Florida, Univ, of Miami Press, Coral 

Gables, FL, 
Naiiman,C,E, 1993, P(ens. In: Flora of North America Editorial C'ommitree, eds, Fl, North 

Amer. 2:1 32-1 35, Oxford Univ, Press, New York and Oxford. 
Radford, A,E,, H.E. Aiii.ns, and C.R. Bi;i.i,, 1968, Manual of the vascular flora of the Caro- 

linas. The Univ. of North Carolina Press, Chapel Hill. 
Small, J. K. 1903- Flora of the southeastern United States. Pubhshed by the author, New 

191,3. Flora of the .southeastern United States, 2nd ed. Published by the 

author, New York, 

1938. Ferns of the southeastern United States. The Science Press Printing 

Co., Lancaster, PA, 

Smith, A.R. and T, Lpmif.dx, 1993- Pteridaceae. In: J,C, Hickman, ed. Thejep.son manual, 
higher plants of California, Pp, 439-1 1 I, Univ, of California Press, Berkeley, 

Stanford, J. W. 1976, Keys to the vascular plants of the Texas Edwards Plateau and atlja- 
cent areas. Published by the author, Brownwood, TX, 

Taylor, R,J, and C,E,S, Taylcjr, 199 i. An annotated list of the ferns, fern allies, gymno- 
sperms and flowering plants ot Oklahoma, 3rd ed. Southeastern Oklahoma State Uni- 
versity, Durant, 

SrDA 18(1): 360, 1998 

Notes 361 


In July of 1997 an unknown plant was collected in Houston and sent to 
Texas A&M for identification. It proved to be Lycianthes asarifolia (Kunth 
& Bouche) Bitter, a member of the Solanaceae native to South America. This 
species has been previously known in North America only from New Or- 
leans, Louisiana (Darwin & Feibelman 1991 )• De Rojas and D'Arcy (1997) 
incorrectly cited that collection as being from Texas. 

This species is recognizable by its prostrate habit, having slender stolons 
that root at every node. The leaves are strongly cordate, entire, and long- 
petioled. Sometimes a second leaf is produced at non-flowering nodes: if 
the primary leaf is relatively large, the second leaf is usually small and of- 
ten stipule-like; when the primary leaf is relatively small, the second is of- 
ten nearly the same size. The flowers are white, rotate, and ca. 1.5 cm broad. 
The yellow anthers are connivent around the style and poricidally dehis- 
cent. The fruit is an edible reddish-orange berry ca. 1.3 cm in diameter, 
closely subtended by the truncate calyx. Plants are self- infertile (Dean 1997). 
A good illustration can be found in the article by De Rojas and D'Arcy and 
images of the Houston plants can be seen at Texas A&M's Bioinformatics 
Working Group Image Gallery page ( 
imaxxsol.htm). Note that the leaves of the Houston plants are nearly all 
rounded apically rather than pointed as in the article illustration. 

A visit to the collection site revealed that this plant has overrun several 
residential yards in Houston, forming a dense, attractive ground cover in 
shaded areas. It is apparently reproducing both vegetatively and sexually, 
for numerous flowers and immature fruits were seen in early December 1 997 
and mature fruits were collected in March 199H. (Darwin and Feibelman 
found no fruit in the Louisiana population and none has since been seen on 
plants grown from cuttings taken from that population.) The authors also 
suspect that propagules are being carried from yard to yard on the equip- 
ment of landscape maintenance companies. The exact time and point of 
introduction is not known, but tlie population has been in existence for three 
or more years. Residents' attempts to control the plant by hand-pulling, 
mowing, herbicide application, and removal of infested sod have proved 
unsuccessful. It appears to be tolerant of Houston's winter weather and suffers 
only partial dieback during the hottest summer days. Should this plant become 
established in nearby Memorial Park (a large, wooded area), it could be nearly 
impossible to eradicate. Herbicide trials were begun in one of the Houston 
yards in March of 1998. After two rounds of tests, no treatment has pro- 

SiDA 18(1): 361. 1998 

362 Sum 18(1) 

vided complere control, and creacmenrs which have weakened the LyciaJithes 
have also damaged the surrounding turf. 

It is possible that this plant could become a noxious weed in southern 
Texas. The USDA Animal and Plant Health Inspection Service is conduct- 
ing a formal risk assessment. (The Louisiana Department of Agriculttn-e 
has considered surveying for the plant in the New Orleans area.) The au- 
thors encourage identification, documentation, and destruction of any new 
colonies subsequently discovered. Should other populations of this plant 
be fotmcl, please contact one of the authors. 

Voucher specimen: U.S.A. TEXAS. Harris Co.: Houston, shaded residential yard of 
502 Lindenwood, with Q^iiercus virginiaiici. Piniis, Stenotaphri/in sec;inclatum , and Trachelosperiiiinn 
m moist sandy loam, 6 Dec 1997, Mary Kelchersnl 1 20697 ^A (BH, BRIT, MO, NO, NY, 

We thank Michael Nee (NY) for his assistance in identifying this plant. 
— Moniq/fe Dubrule Reed, Biology Departvient, Texas A&Al University, College 
Station . TX 77843-523S. U.S.A. and Alary Ketchersid, Agricultural and En- 
vironmental Safety. Texas Agricultural Extension Service. College Station. TX 77845- 
2488. U.S.A. 


Darwin, S.P and T Fiubi' 1 99 1 . Lyda>ithes asarifoih/ (Solanaceac) new to North America. 

Sida 14:605-606. 
Dean, E. 1997. Personal communication. Dr. Dean studies the taxonomy of Lyc/a/zllm at 

the University ot California-Davis. 
Di- RojAS, C.E. and W.G. D'Arcy. 1997. The genus Lycnnithcs (Solanaceac) in Venezuela. 

Ann. Missouri Bot. Ciard. 8-1: 1 67-200. 

SiDA 18(1): 362. 1998 

Notes 363 


Burmeistera Triana (Campanulaceae: Lobelioideae) is a genus of 96 species 
endemic to the Neotropics (Lammers 1998); it was last monographed by 
Wimmer ( 1 943). The genus is closely related to two other Neotropical endemics, 
Centropogon C. Presl and Siphocampyhis Pohl (Pepper et al. 1997; Lammers 
1998). All are large robust plants (herbaceous or suffruticose perennials, 
shrubs, subshrubs, or lianas) with large flowers (corollas averaging 30—60 
mm long or more) borne singly in the axils of the upper leaves or aggre- 
gated into terminal bracteate or foliose (sometimes corymbiform) racemes; 
the tube of the corolla is neither fenestrate nor cleft dorsally and if the lobes 
are dimorphic, it is the two dorsal ones that are the larger. Burnmstera is 
distinguished from its allies by its combination of usually ebracteolate pedicels, 
green or yellow corolla often suffused with maroon or purple, large falcate 
or reflexed dorsal corolla lobes, the wide open orifice of the anther tube, 
baccate often inflated fruit, and oblong to fusiform seeds much longer than 

Most of the species oi Burmeistera are found in montane areas from Costa 
Rica to Ecuador. At the southern limits of this range, two Ecuadorean spe- 
cies extend south into Peru (Stein 1987). At the northern end, a single spe- 
cies is known to occur north of Costa Rica: Burmeistera virescens (Benth.) Benth. 
& Hook, ex Hemsl. It has been reported (McVaugh 1943; Wimmer 1943; 
Nash 1976) only from Guatemala, where it occurs in the departments of 
Alta Verapaz, Baja Verapaz, Huehuetenango, Quezaltenango, San Marcos, 
and Suchitepequez. This is a disjunction from its congeners of nearly 700 
km. That gap has been narrowed, however, by the discovery of this same 
species in central Honduras. This is the first report of any member o^ Burmeistera 
from that nation. 

Voucher specimen: HONDURAS. Depto. Olancho: Parque Nacional laMuralla, trail 
above Visitors' Center, ca. 14 km above La Union, 1 5°()5'N 86°4()'W, in dense high pri- 
mary (2iiercus forest, 1400 m, 27 Oct 1996, P.J.M. Mcun & H. Aiaas 8441 (U). 

The new locality in Honduras is approximately 350 km east of the near- 
est conspecific populations in Guatemala. The 0//«r//j-dominated primary 
forest there was quite rich in epij^hytes, including Pleurothallis tuerckheimii 
Schlecht. (Orchidaceae), Columnea rulyrecaulis Standi. (Gesneriaceae), and Peperomia 
hoffmannii C. DC. (Piperaceae). Undcrstory shrubs included Besleria solanoides 
Kunth (Gesneriaceae), hionnina jerreyrae Taylor (Polygalaceae), Tournefortia 
sp. (Boraginaceae), scandent Sphyrospermum majus Griseb. (Ericaceae), and a 
treelet species oiClethra (Clethraceae). Species found in the herbaceous layer 

SiDA 18(1): 363. 199<S 

364 Si DA 18(1) 

were Psilochilm macrophyllus (Lindl.) Ames (Orchidaceae), Renealmia mexicana 
Klotzsch ex Peterson (Zingiberaceae), Smilacina sp. (Asparagaceae), Spige- 
lia sp. (Spigeliaceae), the saprophyte Gymnosiphon s/iaveokns (Karst.) Urb. 
(Burmanniaceae), and the root-parasite Monotropa uniflora L. (Monotropaceae). 
Adjacent patches of secondary forest contained small trees oiSaurania veraguensis 
Seem. (Actinidiaceae), shrubs oiTriumfetta specwsa Seem. (Tiliaceae), and the 
herbs Carina tuerckhetnni Kraenzl. (Cannaceae) and Hydrocotyle viexicana Cham. 
c& Schlecht. (Apiaceae). 

— Thomas G. Lammers, Department oj Botany, Field Aiuse/rm of Natural His- 
tory, Chicago, IL 60603-2496, U.S.A.; and P.J. M. Maas. Department of Plant 
Ecology and Evolutionary Biology, Herbarium Division. WillemC. van U nnikgebouw, 
Heidelberglaan 2, 3584 CS Utrecht. Netherlands. 


Lammers, T.G. 1998. Review ot the Neotropical endemics Burmeistem, Centropogon^ and 

Siphocampylus (Campanulaceae: Lobelioideae), with description of eighteen new species 

and a new section. Brittonia 50:23.^—262. 
McVaugh, R. I9'13. Campanulaceae (Lobelioideae). N. Amer. Fl. 32A:1-134. 
Nash, D.L. 1976. Campanulaceae. In; Flora of Guatemala, part IX, no. 4. ['ield Mus. Nat. 

Hist., Bot. Sen 24:276-431. 
Pnppi-R, A.S.-R., M.H.G. Gu.stafsson, and V.A. Albert. 1997. Molecular systematics of 

Neotropical Lobelioideae (Campanulaceae), with emphasis on B/innasliri/. Centwpogon. 

and SiphiKampyliis. and the utility of friut and floral characters in lobeHoid classification 

{abstract], Amer. J, Bot. 84(6, Suppl,):222, 
Stein, B.A. 1987. Synopsis of the genus 6«rwe/.f/6'rrf (Campanulaceae: Lobelioideae) in Peru. 

Ann. Missouri Bot. Gard. 74:494-496. 
WiMMHR, F.E. 1943. Campanulaceae-Lobelioideae, I. Teil. In; R. Mansfeld, ed. Das Pflanzenrcich, 

IV.276b. Verlag Wiliielm Engelmann, Leipzig. 

SiDA 18(1): 364. 1998 

NoTi-s 365 


Prior to this report the occurrence of Triraphis and T. mollis R. Br. (purple 
plumegrass) in naturahzed populations was not documented in the United 
States (Chase 1951; Correll & Johnston 1970; Gould 1975; Hatch et al. 
1990; Kartesz 1994; Jones et al. 1997). Specimens of this taxon were first 
collected in naturalized populations by William Godwin on 15 Mar 1993 
in Dimmit County. Since the original U.S. collection, William E. Fox III 
collected the species in the same area on 1 May 1996. 

Cultivated specimens collected from an experiment station in Biloxi, 
Mississippi were located in TAES. However, specimens have not been lo- 
cated that were from escaped or naturalized populations. TROPICOS has 
reported this species as being present in the U.S. because of a listing by the 
U.S. Department of Agriculture in their "National List of Scientific Plant 
Names" (U.S.D.A. 1982). In that U.S.D.A. publication, the distribution 
was not listed because the plant species was not naturalized. 

The species was identified to genus using Clayton and Renvoize (1986) 
and to species using Simon (1993). The identification was then verified using 
the Tracy Herbarium (TAES) plant specimens from Australia and specimens 
from U.S. National Herbarium (US). Lazarides (1970) reported the grass 
to be grazed in Australia, but stated that it appeared to be less valuable 
than many short-lived perennials. Clayton and Renvoize (1986) reported 
the genus to be allied with Neyraudia because of gross morphology and the 
slender microhairs. 

Superficially, T. mollis resembles Aristida. However, upon examination 
of the spikelets with a dissecting microscope the following observations become 
apparent. Spikelets have several florets with the reduced floret apical. Also 
the lemmas are 3-veined, 3-awned. The lemma awns are extensions of the 
3-veins of the lemmas. In T. mollis, the central lemma awn originates from 
the lemma apex and lateral awns below and on both sides of the central 
awn, whereas in Aristida the lemma awns originate from a column or the 
lemma apex and branch from a common point of origin. 

Triraphis mollis R.Br., Prodr. El. Nov. Holl. 185. 1810. (Fig. 1). 

Caespitose perennial. Culms to 50 cm tall, erect to ascending, unbranched; 
nodes glabrous, purplish black; internodes solid. Leaves mostly basal; sheaths 
glabrous, rounded, margins free; auricles minute; ligules a fringe of hairs 
about 1 mm long; blades 6—17 cm long, 1—2 mm wide, flat becoming in- 
volute, glabrous. Panicles 7—20 cm long, 8—15 mm wide (excluding awns), 
contracted, somewhat interrupted, purplish tinged. Spikelets 4—12 mm long. 

Sum 18(1): 365. 1998 


Si DA 18(1: 

Fit;. 1. Triraphis mollis. A. habit; B. spikclet, |;lumes separated trom Horets; C. Horct, palca 
view with lemma margin and the associated hairs; and D. ligule. 

SiDA 18(1): 366, 1998 

Notes 367 

crowded, pedicellate, laterally compressed, disarticulation above glumes and 
between florets; florets 4—10, callus bearded, reduced florets 1-2 and api- 
cal. Glumes subequal (not consistent in which glume is longer), 1 -veined, 
hyaline, shorter than spikelets, awnless; first glumes 3.5—5 mm long; sec- 
ond glumes 4—5.5 mm long; lowermost lemmas 4—5 mm long, 3-veined 
and 3-awned, deeply bifid with teeth to 1 mm long, membranous, median 
vein awns 6—9 mm long from between teeth of bifid apex; lateral vein awns 
5.5—7 mm long, divergent; lateral veins with long pilose hairs (hairs occa- 
sionally on the lemma body); lowermost paleas 2—2.5 mm long, 2-keeled, 
hyaline, glabrous to scaberulous. Stamens 3; anthers 0.2—0.3 mm long, yellow. 
Collection localities were characterized by deep sandy soils of the Brystal 
Fine Sandy Loam (Stevens & Arriaga 1985). Historically, well drained sandy 
soils derived from Carrizo Sandstone were called Norfolk Fine Sand (Carter 
1931). These soils are derived from parent material in the Carrizo Sands 
Formation. Carrizo Sand has been shown to have hygroscopic properties that 
ameliorate the effects of a xeric climate and consequently influence its plant 
formation (McBryde 1933). 

Specimens examined: AUSTRALIA. New South Wales: Sydney; 18??,_/. Maiden s.n. 
(TAES). Northern Territory: Simpson Desert, ca 15 km N of Andado Homestead (250 
km SE of Alice Sprmgs), f 1 Jul \96^J.Weher 9^« (TAES). Queensland-Warrego Dis- 
trict: Charleville, cemetery, Oct-Nov 1945, A'l, Clemens s.n. (TAES). 

UNITED STATES. Texas-Dimmit Co.: 14 mi NW of Carrizo Springs on Cometa Road, 
near sandy dirt road between Cometa and Hwy 277, 15 Mar 1993 W'^ Godwin s.n. (TAES); 
13 mi W of Carrizo Springs, 2 mi W of EM 393 along Hwy 277, then 2 mi N on Cometa 
Road, ak)ng W side of road, 1 May 1996 W.E. Fox III. 548A (TAES). Associated with Pappophor/m. 
Pennisetuni. Set arid. Acacia and Prosopis. 


Financial support was provided in part by Texas Higher Education Co- 
ordinating Board— Advanced Research Program and the Texas Agricultural 
Experiment Station. 

— Stephan L. Hatch, William E. Fox III, and John E. Dawson III, S. M. Tracy 
Herbarium. Department of Rangeland Ecology and Managemmt, Texas A&M University, 
College Station, TX 77843-2126 U.S.A.; and William B. Godwin, Department 
of Entomology, Texas A&M University, College Station, TX 77843-2475 U.S.A. 


Carter, W.T. 1931 . The soils of Texas. Texas Agric. Exp. Sta. Bull. 431 . 
Chase, A. 195 1. Hitchcock's manual of the grasses of the United States, 2nd. ed., U.S.D.A. 
Misc. Publ. No. 200. U.S. Government Printing Office, Washington, D.C. 

SiDA 1H(I): 367. 1998 

368 Si DA 18(1) 

Clavi'ON, W.D. and S.A. Renvoize. 1986. Gener;i graminum, grasses of thf world. Kew 
Bull. Atldit. Scr. XIII. Her Majesty's Stacionary Office, London. 

C()Riu:i.i,, D.S. and M.C. Johnston. I9"(). Manual of the vascular planrs of Texas. Texas 
Research Foundation, Renner. 

Gori.n, F.W. 1975. The grasses of Texas. Texas A&M University Press, College Station. 

Haic:ii,S.L., K.N. Gandhi, and L.E. Browx. 1990. Checklist of the vascular i^lants of Texas. 
Texas Agric. Exp. Sta. Bull. MP- 165 5. 

Jones, S.D., J.K. Wiffe, and P.M. Montgomery. 1997. Vascular plants of Texas, a compre- 
hensive checklist including synonymy, bibliograj^hy and index. University ot Texas Press, 
Austin., J. 1 994. A synonymized checklist of the vascular flora ol the United States, Canada 
and Greenland. Univ. of North Carolina Press, Chapel Hill. 

Lazaridp.s, M. 1970. The grasses of central Australia. Australian Natl. Univ. Press, Canberra. 

McBkydi;, J.B. I9.r3. The vegetation and habitat factors ol tlie (^arrizo Sands. Ecological 
Monogr. .r2l7-297. 

Simon, B.K. 199.>. A key to Australian grasses. Queensland Department of Primary In- 
dustries, Brisbane, Australia. 

Stevens, J, W. and D. Akriaga. 1985. Soil survey of Dimmit and Zavala counties, Texas. 
U.S.D. A.— S.C'.S and Texas Agricultural Experiment Station. 

SiDA 18(1): 368. 1998 




This paper is part of an ongoing study by the author of the vascular plant 
genus Agalinis along the Gulf Coastal Plain. The objective of this note is to 
clarify one of the many nomenclatural difficulties associated with North 
American species of this genus. 

Pennell (19 1 3) described three species of Agalinis from Florida: A. delkatula, 
A. harperi, and A. pinetorum. These species have recently been shown to be 
conspecific (Canne-Hilliker and Kampny 1991). Later, in 1929, Pennell 
reduced A. delicatida to a variety of A. pinetorum as A. pinetorum var. delicatula 
(Pennell) Pennell, considering it to be a weak variety at best and stating in 
his description that it was "Of doubtful rank" (Pennell 1929)- In their study, 
Canne-Hilliker and Kampny (1991) concluded that that all three names 
represented one species and and chose A. harperi for this complex. Kartesz 
( 1 994), although apparently aware of the work of Canne-Hilliker and Kampny, 
chose instead to combine the two species under the name A. pinetorum. 
Although Pennell later recognized the name Agalims pinetorum var. delicatula 
(Pennell) Pennell at the varietal rank, it must still be taken into account for 
nomenclatural purposes because the name A. delicatula has equal priority 
with A. pinetorum (Pennell 1913)- 

The name Agalims harperi and a brief description of the species first ap- 
peared in the Elora of Miami, published in 1913 (Small 1913), in which 
Pennell prepared the treatment oi Agalims. On 13 August 1913, the August 
issue oi Bulletin of the Torrey Botanical Club was published, and in this issue 
Pennell published for the first time the description of A. delicatula and A. 
pinetorum (Pennell 1913). It is clear that Pennell realized that Small's flora 
had appeared (or would appear) before the August 1913 publication of the 
Bulletin, this is because he (p. 426) noted that A, harperi was first described 
in the Elora of Miami. But due to the fact that Small published the Elora of 
Miami himself, I have been unable to determine an exact publication date 
for it. However, in accordance with Art. 31-1 of the Code (Greuter et al. 
1994), A. harperi was effectively published prior to publication of the 13 
August 1913 issue oi Bulletin of the Torrey Botanical Club. The valid publi- 
cation date is fulfilled in two ways in accordance with Articles 29-1 and 
31.1 of the Code: 

1. The preface of Small's Elora of Miami is dated 26 April 1913 (Small 
1 9 1 3); this is the only exact date found in the flora. According to Art. 31.1 
of the Code, in the absence of any other evidence validating the date of 
a work, the earliest date indicating publication is to be accepted as the 

SiDA L8(I): 369. 1998 

370 SiDA 18(1) 

correct publication date. 

2. In a 1997 communication with John F. Reed, Director of the LuEsther 
T. iVIertz Library at the New York Botanical Garden, I was able to verify 
that the publication of Small's Flora of Miaftu did in fact precede the 13 
August 1913 publication date of Bulletin of the Torrey Botanical Cl//b-i\ copy 
of Small's Flora of Miami at the New York Botanical Garden's library is 
date-stamped 21 June 1 91 3. ThLis, publication oi Ai^aTniis harperi in Small's 
Flora of Miami obviously preceded the 13 August 1913 publication of 
A. delicatula and A. pinetonnn in the Bulletin. The valid name for this 
species and its synonyms is as follows: 

Agalinis harperi Penned, in Small, Flora of Miami, 176. 1913. 

A^cilniis ddicatulci Pennell, BliII. Torrey Bo:. Club 40:425. 1913. Agalinis pinetoriiDi var. 
delicatula (Pennell) Pennell, Proc. Acad. Nat. Sci. Philadelphia 81:172-173. 1929. 
Gminlia p/dchella \i\v. dcliailnL/{V{:nnv\\) l^ennell, Acad. Nar. Sci. Philadelphia Monogr. 
1:441. 1935. 
A. pinetorum Pennell, Bull. Torrey Bot. CluLi 40:424-425. 1913. 

Gerardia harperi (PenneW) Pennell, Acad. Nat. Sci. Philadelphia Monogr. 1 : 14 1 . 1935. 
C. piih-bdla (Pennell) Pennell, Acad. Nac. Sci. Philadelphia Monogr. 1:140. 1935. 


I thank John Thieret of Northern Kentucky University for reviewing the 
manuscript and John F. Reed of the LuEsther T. Mertz Library at the New 
York Botanical Garden for his assistance. I also thank Judith Canne-HiUiker 
of the University of Guelph in Ontario, Canada for sharing her unsurpassed 
knowledge of Agalinis with me. — -John F. Hays. Ozark Environmental Con- 
sulting. 3308 Keenes Edge Drive. Columbia. MO 65201. U.S.A. 


CANNii-HiLLiKER, J.M. and CM. Kamp.ny. 1991. Taxonomic .significance of leaf and stem 
anatomy oi Agalinis (Scrophulariaceae) from the U.S.A. and Canada. Canad. J. Bot. 69:1935- 

Greutrr, W., ER. Burdet, W.G. Chaloner, V. Demoulin, D.L. EIawk.swortii, RM. Jorgensen, 
D.E^^, NicOLSON, P.C. Lii.vA, P. Treiiane, and J. McNeil. 1994. International code of bo- 
tanical nomenclature (Tokyo Code) adopted by the Eifreenth International Botanical Congress, 
Yokohama, August-September 1993. Regnum Veg. 131. 

Kartesz, J.T. 1994. A synonymized checklist ot the vascular flora of the United States, 
Canada, and Greenland, 2d ed. Timber Press, Portland. 2 vols. 

Pennei.l, F.W. 1913. Studies in the Agalinanae, a subtribe of the Rhinanthaceae II. Species 
of the Atlantic coastal plain. Bull. Torrey Bot. Club 40:401-439. 

Pi;nnell, EW. 1929. Agalinis aiul allies in North America-ll. Proc. Acad. Natl. Sci. Phila- 
delphia 8 1 : 1 1-249- 

Pennell, E.W. 1 935. The Scrophulariaceae of eastern temperate North America. Acad. Nat. 
Sci. Philadelphia Monogr. 1:119-476. 

Small, J. K. 1913. Elora of Miami. Published by the author. New York. 

SiDA 18(1): 370. 1998 



Biology Library 

University of North Carolina 

Box 5280 Coker Hall 

Chapel mil NC 27399-3280 U.S.A. 

Heinz (Harry) Dietrich Lubrecht, age 88, well known botanical and natural 
history bookseller, antiquarian book expert, appraiser, and former execu- 
tive of the publishing company, Stechert-Hafner, died 12 November 1997, 
at the Columbia Presbyterian Hospital, New York City, after suffering a 
stroke. Harry, as he was affectionately known by friends, colleagues, and 
family, devoted nearly 70 years to the scholarly book trade, particularly in 
botany and natural history. In 1 974 he established Lubrecht & Cramer, Booksellers 
and Publishers, currently located in Port Jervis, NY. His high standards of 
business, steadfast honesty, and breadth of knowledge of botanical litera- 
ture earned him national and international respect as a bookman. With the 
passing of Harry Lubrecht the botanical and bookselling community has 
lost a faithful friend and serious scholar. 

Born 2 December 1908, in Reutlinger, Germany, Harry was the son of 
Adolph August Karl Friederich Wilhelm and Sophie Amalie (Grueninger) 
Lubrecht. His father, an architect, was the son of Lutheran Pastor Adolph 
Lubrecht and Emma Karoline Kuebel. His mother, well-educated and adept 
in three foreign languages, was the daughter of Johann Matthaus Grueninger, 
Professor of Mathematics in the Reutlinger Hochschule, and Sophie Kaemerer. 
When Harry was five, his parents separated. Subsequently, Harry, his mother, 
and two sisters moved to Stuttgart to live in an aunt's apartment home. He 
attended Friedrich Eugens Oberrealschule and completed Hochschule in 
Stuttgart. Throughout his school years Harry was an avid reader. In order 
to ensure enrollment at his school, Harry secured an annual scholarship by 
placing in the top five of his class. 

Upon graduation at age 17, Harry became an apprentice in Brueninger's 
department store as part of his continued schooling. His flare for business 
was well recognized by the store's management and in three years he earned 
the salary of a regular employee. Refusing to join the Nazi party, he was 
compelled to leave Germany. Harry emigrated to the United States where 
his Uncle Christian Grueninger, the mayor of Valhalla, NY, would sponsor 
him. Arriving in America on 15 May 1928, he soon found a job as a grave 

SiDA 18(1): 371. 1998 

372 SiDA 18(1) 

digger ar the local cemetery in Mt. Kensico, NY. When rhis seasonal work 
concluded char fall, Harry looked for employment in New York City. In 
October 192S, G.E. Stechert & Co.,^ well-known international book firm, 
hired him at a weekly salary of $14.00. The stage was thus set for a distin- 
guished, life-long career in the book world, a profession practiced by an- 
cestors on two sides of his family. Some of the Kuebels were medieval scribes, 
the book "publishers" of their time. In the early nineteenth century two 
Lubrecht brothers, Heinrich and August Adolph, separately developed ca- 
reers as booksellers in Heilbronn and Blaubeuren, Germany. In the mid- 
nineteenth century August's son, Karl Theobold, emigrated from Blaubeuren 
to Brooklyn, NY Here he Anglicized his name to Charles T. and founded 
with his cousin the publishing company Haasis and Lubrecht. 

At Stechert Harry advanced from his initial position as billing clerk to 
assistant manager of the Antiquarian Department (April 19.^6). He was a 
natural for this job as he was familiar with German, French, Latin, and Greek. 
His facility for languages assisted him in compiling book catalogs and un- 
derstanding the contents of many scientific books, especially those in Ger- 
man, acquired by the company. Since his immediate boss lacked signifi- 
cant experience in the antiquarian book market, Harry enjoyed the freedom 
and breadth to develop the antiquarian department about which topic he 
read widely. Under his supervision the process of searching for and supply- 
ing out-of-print books was significantly accelerated (Anonymous 1952). Among 
Harry's promotions were to: Chief of Reference and Customer Service, G.E. 
Stechert & Co. (spring 1945); Division Chief, Antiquarian and Out-of-Print 
Department, Stechert-Hafner, Inc. (fall 195 1 ); Vice-President and Editor, 
Hafner Publishing Co. (January 1955); and Vice President, Macmillan Publishing 
Co.-Hafner Publishing Division (1969). 

After conducting a four-year study on subject publishing, Harry con- 
cluded that few firms published books on botany. Through Harry's efforts 
a new subsidiary called Stechert-Hafner Service Agency was established in 
1960. Botanists and natural historians are fortunate to have had Harry's 

'a brief chronology of this book Hrm provides an historical perspective. Stechert & Wolff, 
founded on 1 September 1872, was renamed Gustav E. Stechert (1876), G.E. Stechert (1897), 
and G.E. Stechert & Co. (1904). On 1 October 19^6, the G.E. Stechert partnership was 
dissolved and Stechert-Hafner, Inc. (SH) was created with its publishing and reprinting 
activities transferred to the newly established Hafner Publishing Co. In 1 96t) Stechert- 
Hafner Service Agency was established as a subsidiary of SH. Crowell Collier Jvlacmillan, 
Inc. (now Macmillan, Inc.) acquired SH and its subsidiaries (1969). Beginning January 
1974, the corporate name, Stechert Macmillan, Inc., became effective. By 1980 Macmillan, 
Inc. was divided into three parts of which Macmillan Publishing Co., Inc. contained the 
division now called Hafner Press. 

SiDA 18(1): 372. 1998 

Bdrk, Harry Lubrecht 


Harry Lubrechr making a presentation at the Botanical Society of America Meeting, His- 
torical Section, held at The Ohio State University, Columbus, 1 1 August 1987. Photo Courtesy 
Ronald L. Stuckey. 

wisdom in developing this subsidiary. This new division served scholarly, 
scientific and botanical societies by selling their publications and distrib- 
uting scientific and botanical books for European publishers on a large scale. 
The Agency distributed publications for the New York Botanical Garden, 
Missouri Botanical Garden, Texas Research Foundation, and the Hunt Bo- 
tanical Library (Anonymous 1972). During the 1960s and 1970s Harry also 
focussed attention on reprinting book classics in botany within the Hafner 
Publishing Co. Among the reprint series issued were the Classica Botanica 
Americana (CBA) titles by such eighteenth and nineteenth century lumi- 
naries as Humphry Marshall, John Torrey & Asa Gray, C. S. Rafinesque, 
Stephen Elliott, Thomas Nuttall, and William Darlington. Joseph Ewan, 
historian of botany, proposed this series and wrote a bibliographical com- 
mentary for each of the reprinted books (Ewan 1967). Ironically, these fac- 
simile reprints, as well as many others produced by Hafner, have become as 
rare as the originals themselves. The rarity of these volumes was due in part 

SiDA 18(1): 373. 1998 

374 SioA 18(1) 

to their limited printing, high demand by scliolars, and the destruction of 
remaining stock when Crowell CoUier Macmillan, Inc. ptirchased Stechert- 
Hafner and its subsidiaries in May 1969. 

After this corporate change Harry chose to direct the PubHshing Divi- 
sion. He could not, however, abandon his interest in and passion for rare 
and out-of-print books. Using a substantial part of his lunch hour, he would 
work in the Antiquarian Department at noon time. Through his voluntary 
work, a veritable labor of love, Harry raised impressive revenues for the firm. 

While at Stechert, Harry especially enjoyed appraising private collec- 
tions for purchase. He was instrumental in acquiring an outstanding mathematics 
library in 1 937, besides many other collections. Representing his company, 
Harry participated in the First Antiquarian Book Fair in the United States, 
sponsored by the Antiquarian Booksellers' Association of America and held 
in New York City (4-9 April 1960). Confident that the fair would be suc- 
cessful, he prepared an attractive catalog (#277) of rare books, the first ever 
issued from the Stechert-Hafner Antiquarian Department (Anonymous 1 960). 
One of his career highlights was the discovery of 1 1 first editions of the 
works of Linnaeus in a Paris bookshop. These books had an additional his- 
torical value since they were annotated by the French botanist and explorer 
Michel Adanson (1 727-1806), noted for his work on the natural classifica- 
tion of plants (Lubrecht 1986). 

Harry compiled numerotis book catalogs covering general topics, Greek 
and Latin authors, and natural history. Although no longer in print, his 
catalog (#353), "Early American Botanical Works with a Miscellany of Other 
Botanical Rarities" (Lubrecht 1967), provides an historical survey of early 
classics in Nortii American botany. This booklet, the result of five years of 
searching (Anonymotis 1 967), contains an introduction (pp. 3—5) by Harry, 
an annotated list of 217 botanical titles offered for sale, and a section (pp. 
55-60), "Reference Tools for Botany," by Joseph Ewan. The Missotiri Bo- 
tanical Garden plans to issue a facsimile reprint of this item as a keepsake 
of the XVI International Botanical (Congress to be held at the Garden in St. 
Louis (August 1999). Through the enthusiastic support of Harry, Hafner 
Publishing Co. published and assisted in the production of "A Short His- 
tory of Botany in the United States," edited by Joseph Ewan (1969). This 
book was distributed at the XI International Botanical Congress in Seattle, 
WA, August/September 1969. 

Harry retired from Stechert Macmillan on 30 November 1973 and then 
briefly compiled catalogs for Jack N. Bartfield, an antiquarian bookseller 
of New York City. In December 1 974 Harry founded his own business, Lubrecht 
& Cramer, Booksellers and Publishers, which was later incorporated (Au- 

SiDA hSd): 371. 199H 

BuRK, Harry Lubrecht 375 

gust 1 983). Joerg Cramer (deceased 4 June 1 985), a German publisher, specialized 
in reprinting classical botanical books, many of which Hafner had distrib- 
uted in the United States. Harry's wife Anne, and later his son Charles, assisted 
in the business, originally carried out from their country home in Forestburgh, 
NY, and dealing predominantly with new, scholarly books, mostly in botany 
and natural history. The company is also a distributor for Koeltz Scientific 
Books, Gustav Fischer Verlag, and J. Cramer. Although his bookselling company 
primarily was concerned with new titles, Harry continued his profound interests 
in rare books and the appraisal of libraries. Among the botanical collec- 
tions that he appraised were those of Carroll W. Dodge, the New York 
Horticultural Society, Emanuel D. and Ann W. Rudolph, William C. Steere, 
and R. Gordon Wasson. The remarkable Rudolph Library of 53,()()() vol- 
umes of botany and natural history books has since been donated to The 
Ohio State University Libraries (Stuessy et al. 1997). 

For more than two decades Harry and Anne Lubrecht featured booksale 
tables at botanical and scientific meetings and regional mycological forays. 
Their service was known not only for the sale of books, typically sold at a 
special discount for meeting and foray attendees, but also for their conge- 
nial and knowledgeable conversations. Harry captivated customers with his 
storehouse of botanical and bibliographical knowledge. He and his wife regularly 
attended the annual meetings of the Council on Botanical and Horticul- 
tural Libraries, Inc., and national and international book fairs. Concerning 
the latter, he attended 49 annual exhibits of the Frankfurt Book Fair in Germany. 

Harry was a modest and caring person, yet he forthrightly and candidly ex- 
pressed his opinions. I remember him also for many reminiscences of his book 
trade career and the botanists with whom he associated. With a rich background 
of experience and strong sense of the book market, Harry successfully intro- 
duced the quality paperback, also known as the soft bound, to the scholarly book 
trade. Among the first in his field, he adopted photo-offset printing as the pro- 
cess for producing reprints. Even when this was not widely practiced, Harry 
encouraged minority hiring and developed a staff fluent in numerous languages. 
He held membership in the American Anriquarian Booksellers' Association, 
American Institute of Biological Sciences (AIBS), Council on Botanical and 
Horticultural Libraries, Inc., International Association for Plant Taxonomy, 
Mycological Society of America, New York Botanical Garden, North Ameri- 
can iVLycological Association, and Old Book Table (NY). Harry became a U.S. 
citizen on 14 December 1936. As a member of the Lutheran Church, he was a 
deacon during the 1960s and early 1970s. Among his honors was a Special Service 
Award from the Mycological Society of America presented at the annual meet- 
ing of the AIBS, San Antonio, TX, August 1991. 

SiDA 18(1): 375. 1998 

376 SiDAl8(l) 

Harry's successful life was shared with his wife Anne (Johanne Marie Ficke) 
whom he married on 2 October 1937. Harry is survived by his wife Anne; 
two sons, Peter Thomas Lubrecht, Sr., a retired teacher of Drama and En- 
glish and presently Artistic Director of Brundage Park Playhouse, of Randolph, 
NJ, and Charles Frederick Lubrecht, co-manager of Lubrecht & Cramer, of 
JMontgomery, NY; three grandsons: Peter T. Lubrecht, Jr., Christopher C. 
Lubrecht, and Bryan Leone. Harry's two sisters, Use and Ruth Lubrecht, 
predeceased him. Funeral services were held 15 November 1997 in St. Peter's 
Lutheran Church, Port Jervis and cremation took place at the H. G. Smith 
Crematory, Stroudsburg, PA. Interment of his ashes will take place at the 
Forestburgh Cemetery. As a tribute to Harry, a memorial program in cel- 
ebration of his life will be held at The New York Botanical Garden in the 
fall 1998. Memorial contributions may be sent to The LuEsther T. Mertz 
Library, New York Botanical Garden, 2()0th Street and Southern Boule- 
vard, Bronx, NY 10458. 


I am grateful for information on Harry that Anne, Charles, and Peter 
Lubrecht provided. Ronald L. Stuckey suggested this biographical essay and 
supplied the photograph. The aforementioned individuals and Rogers McVaugh 
read the manuscript and made constructive suggestions for changes in the 
text. Additional information on Harry was obtained from a feature column 
on bookmen (Cfhernofsky] 1986). 


Anonymous. 19')2. Out-of-Print Departmfnt expands. Stechert-I lafner Book News 6(6):H8. 

1960. First Antiquarian Book Fair in America. Antiquarian Bookman 

25(18):1"'{) 1-1706. 

1967. Early American botany catalog. Stcchert-Halner Book News 22(1 ):6. 

.. 1972. 100 years [ol} serving libraries. Stechert-Halher Book News 26(5):H— It. 

C{iiernotsky], j.L. 1986. Lubrecht spans two eras of bookselling history. AB Bookman's 
Weekly 78(5):.i57-358. 

Ewan, J. (cd.). 1 967. Ckissica Botanica Americana, ppi. 38-4(), In {H. Lubrecht], Early American 
Botanical Works with a Miscellany of Other Botanical Rarities. Stechert-Hafner, Inc., 
New York. 

1969. A Short Mistory of Botany in the United States. Hafner Publishing 

Co., New York and London. 

{Lubrecht, H.]. 1967. Early American Botanical Works with a Miscellany of Other Bo- 
tanical Rarities. Stechert-Flafncr, Inc., New York. 

1986. Peregrinations of a rare-bookdealer in Europe. AB Bookman's Weekly 

78(1 ()):808, 810. 

Stuessy, T.E., R.L. Stuckey, W.L. Boomgaarden, and W.R. Burk. 1997. Botanical libraries 
and herbaria in North America. 2. The Rudolph natural history library and its acquisi- 
tion by The Ohio State University. Taxon 46:643—648. 

Sii).-\ 18(1): 376. 1998 

First report of the genus Burmeistera (Campanulaceae) from Honduras 

Thomas G. Lammers and P.J.M. Maas 


Triraphis mollis (Poaceae: Arundineae) a species reported new to the United States 
Stephan L. Hatch, William E. Fox III, John E. Dawson III, and William B. Godwin 

Priority of the name Agalinis harperi (Scrophulariaceae) over the names Agalinis delicatula 
and Agalinis pinetorum 

John F. Hays 

In Memoriam — Heinz Dietrich Lubrecht (1908-1997), botanical bookseller and friend 

William R. Burk 


Book notices and reviews 192, 236, 240, 246 

Index of new names and new combinations in this issue 

Agave gracilis Garcia-Mend. & E. Martinez, sp. nov. 227 

Corsia purpurata var, wiakabui Takeuchi & Pipoly, var. nov. 164 

Crataegus okanagensis J.B. Phipps & O'Kennon, sp. nov. 178 

Crataegus okennonii J.B. Phipps, sp. nov. 170 

Crataegus series PurpureofructiJ.B. Phipps & O'Kennon, ser. nov. 184 

Crataegus phippsii O'Kennon, sp. nov. 185 

Cybianthus anthuriophyllus Pipoly, sp. nov, 89 

Cybianthus cenepensis Pipoly, sp. nov, 106 

Cybianthus comperuvianus Pipoly, sp. nov, 47 

Cybianthus croatii Pipoly, sp. nov, 71 

Cybianthus flavovirens Pipoly, sp, nov, 134 

Cybianthus fosteri Pipoly, sp. nov. 144 

Cybianthus grandezii Pipoly, sp. nov. 139 

Cybianthus granulosus Pipoly, sp. nov. 133 

Cybianthus guyanensis subsp. pseudoicacoreus (Miq. in Mart.) Pipoly, comb, et stat. nov. 49 

Cybianthus huampamiensis Pipoly, sp. nov. 130 

Cybianthus incognitus Pipoly, sp. nov. 125 

Cybianthus jensonii Pipoly, sp. nov. 142 

Cybianthus kayapii (Lundell) Pipoly, comb. nov. 84 

Cybianthus nestorii Pipoly, sp. nov. 59 

Cybianthus pseudolongifolius Pipoly, sp. nov. 101 

Cybianthus timanae Pipoly, sp. nov. 53 

Cybianthus vasquezii Pipoly, sp. nov. 103 

Mandevilla pringleiJ.K. Williams, sp. nov. 231 

Mandevilla holosericea (Sesse & Mog.) J.K. Williams, comb. nov. 237 

Psychotria osiana Takeuchi & Pipoly, sp. nov. 162 

Ruellia jimukensis Villarreal, sp. nov. 223 

Sporobolus pinetorum Weakley & P.M. Peterson, sp, nov. 258 

Trachypogon mayaensis Wipff & S.D. Jones, sp. nov. 242 

Xanthostemon fruticosus Peter G. Wilson & L. Co, sp. nov. 283 

Seasonal changes in concentration and distribution ot heavy metals in creosotebush, 
Larrea tridentata (Zygophyllaceae), tissues in the El Paso, TX/'Ciudad Juarez, Mexico area 
William P. Mackay, Richard Mena, Nicholas E. Pingitorejr., Keith Redelzke. 
C. Edward Freeman. Harold Newman. John Gardea. and Hector Navarro 

New plant records for Dominica, Lesser Antilles 

Steven K. Hill and Arlington James 


A new adder's-tongue (Ophioglossuni: Ophioglossaceae) for North America 
James C. Zech, Patricia R. Manning, and Warren Herb Wagner Jr. 


Notes on the llora of Texas with additions and other significant records 

Larr}< E. Brown and Stuart J. Marcus 


Cyperus fuscus ((^yperaceae), new to Missouri and Nevada, with comments 

on its occurrence in North America 

PaulM. McKenzie. Brad Jacobs. Charles T. Biyson. Gordon C. Tucker, and Richard Carter 


The use of animal-dispersed seeds and fruits in forensic botany 

Barney L. Lipscomb and George M. Diggs, Jr. 



Glaucium corniculatum (Papaveraceae) in Texas 

Zoe Kirkpatnck andJ.K. Williams 


Hovenia dulcis (Rhamnaceae) naturalized in central Texas 
Douglas H. Goldman 


Habranthus tubispathus (Liliaceae) new to the flora of Alabama 
Alrin R. Diamond, Jr., Charles P. Chapman, and Jim Bmmmett 

Plantago coronopus (Plantaginaceae) new to Texas 

Robert J. O'Kennon. George M Diggs, Jr.. and Ro)iald K. Hoggard 


Pteris vittata (Pteridaceae), a new fern for Texas 
Jack W. Stanford and George M. Diggs, Jr. 

Lycianthes asarifolia (Solanaceae) new and weedy in Texas 

Morn que Dubnile Reed and Mary Ketchersid 


(continued on inside back cover) 

ISSN 0036-1488 


Arnoglossuni album (Asteraccac): New s|X'cies Ironi iiorilicrn Florida 
l.iinin C. Aiidei'so}] 


Sc\-en new nonienciauiral coiiihinatioiis and a new name in Packera fAsieraceae: Senecioneae) 
Dchni K. 1 rack and Theodore M. Barkley 

Ardisia niambiensis (Mvrsinaeeae). a new species of Ardisia subgenus Ardisia from ihe ('lioeo 

I'ioristic i^r()\-ince of (iolonibia 

John 1^ Pipoh' 111 (iiid AIniro (J);^olh} P. 


Clusia niambiensis (Ckisiaceae), a new species from the Choco Florisiic Province of Colonihia ami 


John J. Pipoly HI, Alvam Cooallo I' . aiid Maria Sofia Gonzalez 


New species and nomenclatural notes in Clusia (Clusiaceae) from Andean Colomliia and \'ene/uela 

John J. Pipoly HI and Alvaro CogoUuF 


Dos nuevas adiciones a la orquideoflora iMexicana 

Adolfo Kspejo Sema, Ana Rosa Lopez^Femm, Javier Carcia Cruz, Rolando Jimenez Machorroy Luis 

Sanchez Saldana 


Seedling development in species of Chamaesyce (Euphorbun eae) with erect growth habits 
U John Hayden and Olga Troyanskaya 

4 I') 

A revision of the genus Ardisia subgenus Graphardisia (Myrsinaceae) 
John J, Pipoly III and Jon M. Ricketson 


Una nueva especie de Polianthes (Agavaceae) del estado de Oaxaca, Mexico 
Eloy Solano Camacho andAbisai Carcia-Mendoza 


Pollen morphology of the genus Echinopcpon (Cuciirbitaceae) 

Concepcion Rodriguez J. and Rodolfo Palacios-Chdvez 


Variation in the Berlandiera pumila (Asteraceae) complex „ 

G.L. Nesom andB.L. Turner 


New names and combinations in neotropical Myrsinac eae 
J()h)iJ. Pipoly HI and Jon M. Ricketson 


A new species of filmy fern (Hymenophyllaceae: Pteritinphyta) from South India 
C. Ahdiil Ilameed and P V. Madhusoodanan 


A second species of Oritropliium fAsteraceae: Astereai) from Mevico 

(Iiiy L. Nesom 

Unic|Lieness of the endangeied [joricia Semaphore Cacuis (Opuntia cofallitola) 

Ikniiel F. Austin. Darid .\l. Biiininoer. and Donald J. I'inkava 

Ariuinsas Carex (CAperaceae): A lirieflv annoiated list 

Philip E. Hyatt 


(continued on thick corer) 




NuMni'R 2 
Dhcfmbhr, 1998 



I'oi \i)i:i) li^ 

Lloyd H, Shinnei 


Wm. !■. Mahler 
PulilisluT 19^1-1992 

— - vf 

l^ai"nL'\" I.. Lipscomb 


Hoianical Kcscaaii liisiiiuic ot Tcxa.s, Inc. 

Sn9 Pecan Slivct 

I'ort \\'o\-\h. Texas 7(il02- H)()(), liSA 

817 332- Mil sr 332-a 112 FAX 

I'.leclfoiiic mail: sida@hril.()i;^ 

lome page al the IRL: http: \\\\ w.hnl.orj^ siJa 

John W, Thieret 

Associate hxlitor 

Hiol(\L;ical Sciences Dept. 

Northern Kenluckv l'ni\ersily 

limhlanJ llemhis. Keiitiick\- i KPfi, I'SA 

Prof. Dr. Irli.x Llamas 

Coiitrihulint^ Sjxmish I-^iilor 

Dplo. de Botfmica, L'acullad de Hioloi^i 

llni\ersicLid de Leon 

L>2 i()~l Leon. Spain 

(iiiuk'LiK's lor (."iiiiii"ihiiiiii> aa' avaihibk' upon iwjlk'sI 

• ind 1)11 ihi- inside Ixick cover ot the la-it issue ol] xohinie. 

SuhstTiption per year: S_!t. iiuli\RliiaL Si^. ISA liisiiuilion.s, J>sl), ( )ulsid(.- kSA; 

[lunilxT.s issued twice a \ear 


•'SlDA. (X)NTK11UTI()NS TO BOTAX^'. \oliinie IS, Xiimher 2, jiai^es r^-(^M) 

eopMi^i^hi i')')s 

i5olanieal Kese.uxii liisiiiuie ol Te.\as. Iiic 

Printed in tin- I nitetl States ol Anierua 

ISSX (11)3(1- 1 |S8 



Department of Biological Science 

Florida State University 

Tallahassee, FL 32306-4370, U.S.A. 


The new species Arnoglossum album is described. Its phyllaries have prominently winged 
keels — a feature shared with ^. diversifolium, A. floridanum, A. plantagineum, andy4. sulcatum. 
Comparisons among members of this closely knit assemblage include morphology, phe- 
nology, ecological setting, and range, and a key is provided to distinguish the species. 


Sc describe una especie nueva Arnoglossum album. Sus filarios tienen qtiillas aladas 
prominentcs — una caracteristica que comparte con A. diversifolium , A. floridanum, A. 
plantagineum, y A. sulcatum. Las comparaciones entre miembros de este conjunto fuertemente 
reticulado incluye morfologia, fenologia, caracteristicas ecologicas y areal. Se ofrece una 
clave para diferenciar las especies. 

Species of the x.w?dA?i^\wd\di Arnoglossum, commonly called "Indian Plantains," 
were formerly placed in the heterogeneous '^Cacalia" (Krai & Godfrey 1958; 
Cronquist 1978; Pippen 1978), but, because Cacalia is now typified by a 
species of the senecionoid European genus '' Adenostyles' and because ge- 
neric circumscriptions are now more narrowly drawn, Arnoglossum is con- 
sidered distinct by many recent authors (Robinson 1974, 1980; Jeffrey 1979, 
1992; Godfrey &Wooten 1981; Wetter 1983; Funk 1985; Brummitt 1992). 
Arnoglossum has three species with extensive ranges in the eastern United 
States \A. atrip licifolium (L.) H. Robins, andyl. reniforme {Wook.) H. Robins.] 
or southeastern United States [A. ovatum (Walt.) H. Robins.] and one from 
the central part of the country \A. plantagineum Raf ] . The remaining three 
species have more limited ranges, one as a Florida endemic [A. floridanum 
(A. Gray) H. Robins.] and the others with ranges barely extending from 
Florida into neighboring states [A. diversifolium (Torn & A. Gray) H. Robins, 
andyl. sulcatum (Fern.) Fi. Robins.] . The latter four species {i.e., A. plantagineum 
and "the Florida group") are similar in having phyllaries with keels that are 
strongly winged. The new species described below is now added to this assemblage 
with winged phyllaries. . ■ - • -: 

SiDA 18(2): 377-384. 1998 

378 SiDA 18(2) 


Fresh and dried materials were processed as described by Anderson (1994). 
Heads from at least five different collections per species were dissected and 
measured for involucral and floral quantitative data. Living plants of the 
five species under study were pro]:)agated from seed or as transplants in a 
greenhouse or common garden in Tallahassee, Florida. Vouchers of seed- 
lings, plants at various stages of development, and all cited specimens are 
deposited at FSU unless noted otherwise. 


Arnoglossum album L.C. Anderson, sp. nov. (Figs. 1, 6-9). Tyei,: U.S.A. 
FLORIDA. Bay Co.: locally common in acidic, poorly drained .sandy soil of open, 
wet savanna with Sarracenia flava, Rhynchospord, Rhtxici, and Polygala spp. just E of 
Rte 77 on N side of Southport, T2S, R14W, NElAl of SEl/4 Sec 21, 6 Jun 1995, 
L.C. Amkrmi 75555 (HOLOTYPE: NY!; ISOTYPES: BRIT! FSU! MO! US!). 

Ilerbac perennes plertimqLie 8—1 dm aka. Folia basalia 20—63 cm lon^'a, 5—16 cm lata, 
longe petiolata, laminis late ovatis vel anguste oblongo-lanceolatis, verms lateralibus principalibus 
per 2—4 cm supra basin laminae ad venam mediam arete parallelis dein divergentibus. Folia 
caulina pauca, 7-26 cm longa, petiolata, serrata, foliis superioribus reductis sessilibusque. 
C^apitula in cymis corymbosis disposita, involucris 10—1 3.4 mm longis, piiyllariis albis carinis 
promincnter alatis. CoroUae albae, 9—10.5 mm longae. 

Essentially glabrous cespitose herbs. Stems (5-)8-l()(-l 0.5) dm tall, light 
green, strongly ridged. Basal (radical) leaves (2()-)3()-55(-63) cm long overall, 
5-10(-l6) cm wide, long petiolate, (1()-)1 5-18(-3l) cm, blades broadly 
ovate to narrowly oblong-lanceolate, margins entire, shallowly sinuate, or 
rarely serrulate-denticulate, bases attenuate, apices rounded, mucronulate, 
basally disposed lateral veins appressed to the midrib 2-4 cm then abruptly 
extending toward the leaf margins; cauline leaves few, blades (7-)12— 1 8(— 
26) cm long overall, (2. 5-)4-5(-6) cm wide, petioles (I-)4-5(-l 2) cm long, 
ovate, bases cuneate, apices acuminate, serrately toothed, upper leaves re- 
duced in size and sessile. Inflorescence compound, freely branched, ± flat- 
topped, with heads crowded in corymbose cymes. Heads 5 -flowered; in- 
volucres 1 {)— 1 2.5(— 1 3.4) mm long, cylindric, phyllaries 5, chalky white, 
margins hyaline, keels prominently winged (the wings rising 1 — 1.5(— 2.0) 
mm from the phyllaries with margins sinuate to erose and reduced apically). 
Receptacles with short central cusp. Corollas white, rarely tinged with pink, 
9—10.3 mm long, veins 10, tube 5-6.4 mm, throat 0.2—1 .0 mm, lobes 2.9- 
3.8 mm, lanceolate; anthers 2.5—2.6 mm long, collars cylindrical; styles 1 1 .8— 
12.5 mm long, style branches 1.5-1.7 mm long, stylopodium immersed 
in nectary; achenes glabrous, 4.2—5 mm long, cylindric to narrowly clav- 
ate, carpopodium cells erect, pappus white, 6-7 mm long, n = 25. 

Di.strihution. — Endemic to Bay and Gulf counties of Florida in poorly 

Andrrson, a new species of Arnoglossum 


Figs. 1—24. Representative leaves of selected Arnoglossum species, all at 1/6 full size. Figs. 
1-5. Radical leaves showing major venation. Fig. \ . A. album with appresscd lateral veins, 
Anderson 15600, Bay Co., FL. Fig. 2. A. divcrsifoliunu Anderson 13414, Levy Co., FL. Fig. 
3. A. flortdanurn, Anderson 72/65, Taylor Co., FL. Fig. A. A. plantagineum, Anderson 12166, 
Riley Co., KS. Fig. 5. A. sulcatum, Anderson 16035, Santa Rosa Co., FL. Figs. 6-24. Out- 
lines of cauline leaves (major venation similar to respective radical leaves). Figs. 6—9. A. 
album. Fig. 6. Anderson 12137, Cuh Co., FL. Figs. 7-8. Anderson 15600, Bay Co., FL. 
Fig. ^. Anderson 13432, Gulf Co., FL. Figs. \{)-\5-A. diversifolium. Fig. \'^. Anderson 13475, 
Levy Co., FL. Figs. 11, 13. Anderson 13478, Putnam Co., FL. Fig'^ 12. McDamel 9059, 
Houston Co., AL. Figs. 14—15. A. floridanum. Fig. 14. Anderson 12160, Taylor Co., FL. 
Figs. I'b-lG . Anderson 12077, Clay Co., FL. Figs. 17-20. A. plantagineum. Fig. 17. Mehrhoff 
12299 (CONN), Bruce Co., Ontario Prov., Canada. Fig. 18. Anderson 11991, Loundes 
Co., AL. Fig. 1 9. Bryson 9861, Oktibbeha Co., MS. Fig. 20. Anderson 11968, Tangipohoa 
Par., LA. Figs. 1\ -24. A. sulcatum. Fig. 2\. McDamel 7978, Geneva Co., AL. Fig. 22. Anderson 
12342, Leon Co., FL. Fig. 23. Anderson 16058, Walton Co., FL. Fig. 24. Anderson 16057, 
Walton Co., FL. 

380 SinAl8(2) 

drained, acidic (pH 5-6), loamy sands (with 0.35-1.7% organic matter) 
of wet savannas, open pinewoods, and most frequently the ecotone between 
the two, often in close association with several of the following: Aletris liitea, 
Aristida heyrichiana, Asclepias longifolia, A. michauxii, A. viridula. Aster 
eryngiifolius, Cleistes bifaria, Drosera capillaris, Eriocaidon compressum, Hy- 
pericum gymnanthum, Lachnanthes caroliniana, Lachnocauion anceps, Lo- 
heliaflorid^ina, Ludwigia linifolia, L. microcarpa, L. virgata, Murshallia tenuijolia, 
Oxypolis greenmanii, Panicum rigididum, Physostegia godfreyi, Platanthera 
nivea, Polygala balduinii, P. cruciata, P. ramosa, Rhexia lutea, Rhynchospora 
ciliaris, R. curtisii, R. fascicularis^ R. filifolia, R. globularis, R. oligantha, R. 
pusilla, R. rari flora, R. wrightiana., Rudbeckia graminifolia, Ruellia noctiflora, 
Sabatia bartramii, S. campanidata^ Sarracenia flava, Scutellaria floridana, 
Tofieldiaracemosa, Verbesina chapmanii, Xyris ambigua, X. baldwiniana, and 
X. elliotii. The woody associates (less frec[uent) include: Cyrilla racemiflora, 
Hypericum cistifolium, H. exile, H. reductum. Ilex coriacea, I. vomitoria. Magnolia 
virginiana, Myrica cerifera, Nyssa ursimi, and Pinus elliottii. Flowers June to 

Representative specimens examined. U.S.A. FLORIDA. Bay Co.: rvpc locality, /..C/l;/('/i:';w;^ 
/ 552.9 (FSU), 20 Jim 1995, L.C. Anderson 15600 (I'SU, MO, NY), 20 (un 1995, L.C. 
Anderson 1 5601 (FSU), L. C. Anderson 15708 (FSU); S side County Rd 2300, 1 .9 mi W of 
Rte 77. 1 .5 air mi NW of Southport, 1 3 Jim 1 995, L.C. Anderson /5577(FSU); 0.5 mi E 
of Butnt Mills C;rcek bridge on S side of Rte 388, 1 .9 mi W of Rte 77, 1 .5 air mi WNW 
ofSouthporr, 13 jun 1995\ L.C. Anderson I558I (FSU). Gulf Co.: 5.3 miSofRte22on 
E side of Daniels Rd near Wetappo Creek, ca. 5.5 air mi WSW of Wewa- hitchka, L.C. 
Anderson 1 1642 (FSU), L. C. Anderson 1 167.3 (2), L.C. Anderson 1 1934 (FSU), L.C. Anderson 
/2/37(BRi'F FSU, CiH, MO, NY, US), L.C. Anderson 121.38 (FSU), L.C. Anderson 1.3421 
(FSU), 5.2 mi S of Rte 22 on W side Daniels Rd, L.C. Anderson 11950 (FSU), L. C Anderson 
12171 (FSU). 


The Indian plantains with winged phyllaries form a closely related alli- 
ance of species. They differ from each other in several floral features as summarized 
in Table 1 . The longest involucres occur in A. floridanum, and the shortest 
in ^. sulcatum.. Phyllaries mA. album have prominent wings proximally 
that become highly reduced apically, whereas in all other species the wings 
are shorter in height but ± equally cieveloped for the length of the bract 
(often broadest apically). Arnoglossum album also has the whitest phyllar- 
ies; in the other species they are more greenish-white. 

Corollas oiA. album are white, rarely tinged with pink; in the other species 
they are white, light lavender, or ochroleucous (more greenish-white or yellowish- 
white in A. floridanum and sometimes more pinkish in A. diversifolium) . 
Corollas are generally longest in A. floridanum and shortest in A. sulcatum, 
but not as short as reported by Pippen (1978) and Cronquist (1980). In all 
species the corolla tubes are relativelv long and the throats below the lobes 

Anderson, A new species of Arnoglossum 381 

Taiu I 1 . Size floral features (in mm) of "winded" Arnoglossum species. 


Involucre length 

C^orolla length 

Tube width 

Lobe length 

A. album 





A. diversifolium 
A. floridanum 





A. plantagmeum 
A. sulcatum 






very short. The tube widths are greatest in A. alburn and most slender in A. 
sulcatum. Carpopodial cells of the achenes in A. album are generally erect, 
whereas they are procumbent in A. plantagineum. Cell shape varies some- 
what within a given population because of differing stages of achene matu- 
rity. Robinson and Brettell (1973) suggested carpopodial structure was an 
excellent character for distinguishing genera in the Senecioneae; they re- 
corded procumbent carpopodial cells iox Mesadenia (i.e., Arnoglossum) .^&\ter 
( 1 983) found considerable variation in this and other micromorphological 
characters and questioned their value as generic markers. 

Vegetatively, yi . album looks most X'^lq A. plantagineum, and that species 
is probably its closest relative. The two differ in seedling morphology. Cotyledons 
ofyl. album are 1-1.5 cm long and 9-12 mm wide (± orbicular), whereas 
they are 3—4.5 cm long and 8—9 mm wide (spatulate-oblanceolate) in ^. 

The five species differ more in leaf morphology than they do in floral 
features (Figs. 1—24). Basal leaves oi A. album have distinctive venation. 
The principal lateral veins are appressed and closely parallel the midvein 
into the blade for a few centimeters and then spread abruptly toward the 
leaf margins (Fig. 1). This feature of appressed lateral veins is also found in 
the related Yermo xanthocephalus ofWyoming. In the oxhei Arnoglossum species, 
the lateral veins spread immediately at the base of the blade (Figs. 2-5). 

Relative length of the petiole on radical leaves is often used in keys to 
the species (Krai & Godfrey 1958; Pippen 1978; Cronquist 1980). This 
feature is correlated to habitat preferences of die species. Generally, A. diversifolium 
andy4. sulcatum grow in deep shade, and their radical leaves have long peti- 
oles (Figs. 2, 5). Arnoglossum floridanum usually grows in full sun, and its 
radical leaves have short petioles (Fig. 3). Petiole lengths are more variable 
in radical leaves of /I. album ^ind A. plantagineum. These two species often 
grow in savannas or prairies. Their petiole lengths are related to the density 
of grass and other vegetation near the plants; plants from the same seed 
source that were grown in different shade or vegetation densities in my garden 
produced short-petioled leaves in sunnier or less crowded situations and 
longer petioles in shaded or crowded conditions. 

382 SmA 18(2) 

Cauline leaves of A. diversifolium (Figs. 1 0-1 3) are periolate and basally 
truncated or cordate; they are petiolate and rounded to broadly cuneate in 
A. album (Figs. 6—9), A. floridanum (Figs. 14—16), -^nd A. piantagineum 
(Figs. 17-20), whereas they are ± sessile and more narrowly cuneate in A. 
sulcatum (Figs. 21-24). Cauline leaves of^. diversifolium are deltoid-hastate 
in oudine and dentately toothed. They tend to be ovate, serrately lobed, 
and acute in A. album; ovate-oblong, crenulate, and obtuse or rouncied in 
A. floridanum\ ovate to lanceolate or narrowly spatulate, entire or serru- 
late, and acute in^l. plantagineum\ and ovate to rhombic or narrowly lan- 
ceolate, sparsely serrate-dentate, and acute to obtuse in A. sulcatum. 

Arnoglossum plantagineum has the greatest geographical range among the 
five species (east Texas, Louisiana, and the black soil belt of Alabama north 
through Kansas and Ohio to southern Minnesota, Michigan, and Ontario, 
Canada); it occurs mainly in calcareous, tall-grass prairie (Pippen & Chapman 
1986), but minor habitats include glades, fens, and pine-oak woodlands. 
Arnoglossum floridanum occurs in well-drained sands in open pine-scrub 
oak, dry flatwoods, and old fielcis from Duval and Madison through pen- 
insular Florida south to Highlands and Manatee counties. 

The other three species occur mainly in the Florida panhandle. Arnoglossum 
diversifolium occurs in river swamps and wet hammocks and is the most 
wide-ranging of these three; it has a few populations in southern Alabama 
and southwestern Ceorgia, ranges fi-om Walton County to Leon County 
in the panhandle, and has disjunct populations in Putnam, Volusia, and 
Levy counties, Florida. Arnoglossum sulcatum occupies shaded acid bogs or 
swamps; it occurs sporadically in extreme southern Alabama and south- 
western Georgia and from F^scambia to Leon Counties of the Florida pan- 
handle. Arnoglossum album is restricted to Bay and Gulf Counties of the 
Florida panhandle; the only other species of the group that occurs in those 
counties is A. sulcatum, but it has different habitat preferences and 

Flowering time in Arnoglossum plantagineum is clinal from south to north; 
the plants blooming in late April in southern Louisiana to early August in 
Minnesota and Michigan (l^ippen & Chapman 1986). A reverse dine ex- 
ists for A. diversifolium; it blooms in the Florida panhandle from May to 
early July (same period fox A. floridanum) , whereas, to the south, in Levy 
County (where it is sympatric with Hasteola robertiorum) A. diversifolium 
blooms from mid-August through September. Arnoglossum album blooms 
from June to mid-July, and A. sulcatum is the latest to bloom (September 
to October). 

The new species, A. album, is distinguished from all of its immediate 
relatives by the following features: it is geographically isolated from all other 
species except A. sulcatum, and it is totally separated from A. sulcatum by 

Andkrson, a new species of Arnoglossum 383 

habitat and phenology; basal leaf venation is distinctive; cauline leaves are 
most similar to those of /}. sulcatum, but they are generally narrower and 
sessile in A. sulcatum; its involucres and corollas appear whiter than those 
of the other species; the wings of the phyllaries are higher proximally and 
attenuated and distally, whereas wings are lower in height overall and evenly 
raised along the keel or somewhat higher distally in the others; the phyl- 
lary wings have erose margins distally in A. album, and wing margins are 
entire or sinuate in the other species; and the corolla tubes are wider than 
those in any other species in the group. 


Most keys to species of Arnoglossum include significant habitat and phe- 
nological data. The following key uses only morphological features. 

1. Blades ot radical leaves truncate, ovate to cordate-ovate, sparsely den- 
ticulate; lower cauline leaves deltoid-hastate, dentately toothed A. diversifolium 

1. Blades of radical leaves ovate to ovate-oblong, entire, sinuate or crenate; 
lower cauline leaves ovate, entire, crenulate or serratcly toothed. 
2. Phyllaries with prominently winged keels, wings highest at base, erose; 
radical leaves with lateral veins appressed to midvcins lor 2—4 cm 

then spreading A. album 

2. Phyllaries with keels + evenly winged throughout or highest apically, 
entire or sinuate; radical leaves with lateral veins diverging from base 
of blade. 

3. Involucres and corollas mostly over 1 1 mm and 10 mm long, re- 
spectively; cauline leaves crenulate A. floridanum 

3. Involucres and corollas mostly shorter; leaves entire or sparsely serrulate. 
4. Involucres mostly over 1 mm long, corollas over 9 mm; cauline 

leaves at midstem rounded to petiokue base A. plantagineum 

4. involucres less than 10 mm long, corollas 8-9(-9.6) mm; cauline 

leaves at midstem broadly cuneate and sessile A. sulcatum 

acknoweedc;m ents 

This study was supported in part by a grant from the Council on Re- 
search and Creativity of Florida State University. Richard Pippen graciously 
shared his knowledge of the group. Mark Garland assisted with the Latin 
diagnosis, and Ken Womble helped with the illustrations. Ted M. Barkley, 
R.R. Kowal and A.B. Thistle are thanked for providing critical reviews of 
the manuscript. 


AndI'RSON, L.C. 1994. A revision of Hasteola (Asteraceae) in the New World. Syst. Bot. 

Brummiti , R.K. 1992. Vascular plant families and genera. Kew: Royal Botanic Gardens. 
Cronqulsi , A. 1978. Compositae, tribe Senecioneae. N. Amer. Flora, sen II, 10:14-16. 
. 1980. Vascular plants of the southeastern United States. Vol. 1 . Asteraceae. Univ. 

384 SinAl8(2) 

of North Carolina Press, Chapel Hill. 

Funk, V.A. 1985. Cladistics and generic concepts in the Compositae. Taxon 34:72-80. 

GoDi-RKY, R.K., and J.W. WocriEN. I 98 1 . Aqtuuic and wetland plants of the southeastern 
United States. Dicotyledons. Univ. of^ Georgia l^'ress, Athens. 

Jl.i FRiv, C. 1979. Generic and sectional limits in Senecio (C'oniposltae): II. F.vahiation of 
some recent studies. Kew Bull. 34:49-58. 

. 1992. 1 he tribe Senecioneac (Compositae) in the Mascarene Islands with an an- 
notated world check-list of the genera of the tribe. Kew Bull. 74:49-109. 

Kral, R., and R.K. GoDFRl-Y. 1958. Synopsis ol the Florida species of ('iicnliii (Compositae). 
Quart. J. Florida Acad. Sci. 21:193-206. 

PiPPHN, R.W. 1978. Cnailia. N. Amer. Flora, series 11, 10:151-159. 

and K.A. CjiaI'MAN. 1986. Comparison oi morphological characters between re- 
gional habitats of Cacalia plantaginea (Asteraceae). In: G.K. Clanby and P.N. Pemblc, 
eds. Proceedings of the 9rh North American Prairie C'onference. IVi-college University 
C^enter for Environmental Studies, Faijjo, ND. 

Robinson, F1. 1974. Studies in the Senecioneae (Asteraceae). VI. Fhe genus Arnoglossiim. 
Phyrologia 28:294-295. 

. 1980. Arnoglossiim rcniforme (Hook.) li. Robins., comb. nov. Phytologia 46:441. 

and R.D. Brettcll. 1973. Studies in the Senecioneae (Asteraceae). IV. The genera 

Mcsiidoi'uL Syiieik'sis, Miricncalicu Koyamacalia and Sirniciiliti. Phytologia 27:265-276. 

Wlil'll.R, M.A. 1983. Micromorphological characters and generic delimitation of some 
New World Senecioneae (Asteraceae). Brittonia 35:1-22. 



Herbarium, Division of Biology 

Kansas State University 

Manhattan, KS 66506, U.S.A. 

dkwel@ksii. edu 


The following new combinations and new name arc proposed: Packera boLinderi var. 
harfordii (Greenm.) D.K. Trock & T.M. Barkley, comb, nov.; Packera dimorphophylla var. 
intermedia (T.M. Barkley) D.K. Trock & T.M. Barkley, comb, nov.; Packera dimorphophylla 
var. paysoni (T.M. Barkley) D.K. Trock & T.M. Barkley, comb, nov.; Packera pseuda urea 
Nss.flavula (Greene) D.K. Trock & T.M. Barkley, comb, nov.; V-A.cks.x-3,pseudmirea\'\\x. semicordata 
(Mack. & Bush) D.K. Trock &T.M. Barkley, comb, nov.; Packera neomexicanavix. tonmeyi 
(Greene) D.K. Trock & T.M. Barkley, comb, nov.; Packera neomexicana var. mercalfei (Greene) 
D.K. Trock & T.M. Barkley, comb, nov.; and Packera buekii D.K. Trock & T.M. Barkley, 
nom. nov. 


Sc proponen las siguientes nuevas combinaciones y nuevo nombrc: Packera bolanderi 
var. harfordii (Greenm.) D.K. Trock &T.M. Barkley, comb, nov.; Packera dimorphophylla 
var. intermedia (T.M. Barkley) D.K. Trock & T.M. Barkley, comb, nov.; Packera dimorphophylla 
var. paysoni (T.M. Barkley) D.K. Trock & T.M. Barkley, comb, nov.; Packera pseudaurea 
\''ix. fldvula (C irccne) D.K. Irock & T.M. Barkley, comb, nov.; Packera /^ifwd^z/rcd var. semicordata 
(Mack. & Bush) D.K. Trock & T.M. Barkley, comb, nov.; Packera neomexicana var. toumeyi 
(Greene) D.K. Trock & T.M. Barkley, comb, nov.; Packera neomexicana var. mercalfei (Greene) 
D.K. Trock & T.M. Barkley, comb, nov.; y Packera buekii D.K. Trock & T.M. Barkley, 
nom. nov. 

The genus Packera (Asteraceae: Senecioneae) has been known widely as 
the Aureoid group oi Senecio, and the recognition o^ Packera necessitates 
many nomenclatural transfers. Preparation of the treatment o{ Packera for 
the Flora of North America project has drawn our attention to the trans- 
fers proposed here. Type specimens for the basionyms are provided by Barkley 
( 1 978) . The combinations P. pseudaurea var. semicordata and P. dimorphophylla 
var. intermedia are proposed simply for consistency in the use of "varietas" 

'Present address: Botanical Research Institute ofTexas, 509 Pecan Street, Fort Worth, TX 

76102-4060, U.S.A, e-mail: 

SiDA 18(2): 385-387. 1998 

386 SiDA 18(2) 

as the infraspecific rank of choice in Packera The two epithets are other- 
wise available as subspecies. 

Packera bolanderi var. harfordii (Greenm.) D.K. Trock &T.M. Barkley, 

Comb.nov. BASIONYM: Senecio Ijarfordii Gveenm. in PipeT, CA)nrr. U.S. N;itl. Herb. 
1 1 :567. 1906. St'iu'do hoLiiideri var. hiirforflii {(;recnm.) I'.M. Barkley 

Packera dimorphophylla var. intermedia (T.M. Barkley) D.K. Irock & T.M. 
Barkley, comb. nov. By\.SIC)NYi\l: Senecw dimoypliophyllmxAX. intcTiiiedius'Y.M.. 
Barklev. Trans. Kansas Acad. Sci. 65:362. 1 963. Packera dimorphophylla subsp. intermedia 
(T.M.'Barkky) Weber & Love. 

Packera dimorphophylla var. paysoni ( l.M. Barkley) D.K. Trock & l.M. 
Barkley, comb. nov. BA.SIONYM: Seuecio dimorphophyllm\':\\. payu^ii^WW . Barkley, 
Trans. Kansas Acad. Sci. 65:362. 1963. 

Packera pseudaurea var. flavula (Greene) D.K. Trock &T.M. Barkley, comb, 
nov. BaSIONYM: Senecio flamdus i^\\xi:nc, Pittonia 4: 108. 1900. Setiecio pseudaureus 
var. flavula (C^reene) Greenm. Packera pseudaurea suhsp. [lavula (Cirecne) Weber & 

Packera pseudaurea var. semicordata (Mack &C Bush) D.K. Trock & T.M. 
Barkley, comb. nov. BaSIONYM: Senecio semieordatus Mack. & Bush, Rep. 
Missouri Bot. Card. 16:107. 1905. Senecio aureus var. semieordatus (Mack. & Bush) 
(u'eenm. Soiecio pseudaureus viw se)nicordatus (Mack. & Bush) F.M. Barkle\'. 

Packera neomexicana var. toumeyi (Greene) D.K. Trock & f.M. Barkley, 
comb. nov. ^i\S\OW\\\: Senecio toumeyi Gvciinc, Pittonia 3:349. \89S. Senecio neo- 
mexicanus \'AV. /o //;?«')'/ (Greene) l.M. Barkley. 

Packera neomexicana var. metcaltei (Cireene) D.K. Trock & F.M. Barkley, 
comb. nov. BASIONYM: Senecio ?netcalfei CiKcne ex Wooron & Standi., (lonlr. U.S. 
Natl. I lerb. 16:193. 1913. Senecio ueowe.\ica>/us v.w. metcalfei [Greene) I'.M. Barkley. 

The situation with the epithet '\ymbaiarioiJef is complicated. A single- 
headed plant of wet, open sites in western U.S. and adjacent Canada was 
long known as Senecio subnudus DC., a name that was introduced by A. P. 
de Candolle in 1 837. Unfortunately, de Candolle used the epithet "subnudus" 
twice; the other time For an African plant. Heinrich W. Buck (1840) rec- 
ognized the error in his index to the Prodromus and he chose the African 
plant to be the correct Senecio subnudus. He then called the American plant 
Senecio cymbalnrioidesW. Buck, and unfortunately, this n